The influence of temperature on the exudation of xylem sap from detached root systems of rye (Secale cereale) and barley (Hordeum vulgare)

Summary Roots of intact plants of rye and barley which had been growing at 20° were cooled for 12–72 h at 8–14° C while the shoots were kept at 20°. The roots were then excised and placed in solutions at temperatures ranging from 2.5–22.5° C. The rate of exudation of xylem sap and the chemical composition and osmotic potential of the sap were measured and compared with controls which had been kept at 20° C during the pretreatment period. Pre-cooling increased the fluxes of K⁺, Ca²⁺ and H₂PO ₄ ^- into the xylem sap of both species by factors of two to three; the total volume of exudate rose by larger factors. Thus the concentrations of these ions were lower in the sap exuding from cooled roots than in that from controls. Measurements of the osmotic potential of the sap from barley roots indicated that the osmotic driving force in cooled and control roots was similar even though flow in the former was much greater.The enhancement of exudation was shown to be dependent on the duration and the temperature experienced by the roots during pretreatment, and was lost rapidly when roots of intact plants were returned to 20°.Analysis of the temperature coefficients for exudation and Arrhenius plots revealed very distinct changes in the activation energy for exudation above and below a transition temperature. In control plants of barley and rye this temperature was around 10° C, but in cooled roots of rye there was a significant shift in the transition temperature to 5° C. Activation energies for exudation of control and cooled roots above or below the transition temperature were broadly similar, thus pre-cooling roots did not alter the temperature sensitivity of exudation but merely its rate at a given temperature.The results are discussed in relation to active ion transport, membrane fluidity and the resistance of the root to water flow.Abbreviation ABA abscisic acid     

Effect of Potassium, and Abscisic and Indole-3-Acetic Acids, on Maize Root Xylem Exudation and Potassium Efflux

The influence of potassium sulfate, abscisic acid (ABA) and indole-3-acetic acid (IAA) solutions on xylem exudation rate and potassium efflux from the apical cut end of root tips of intact maize (Zea mays L. cv. Dnepropetrovskaya) seedlings was studied. Foliar application of 5 mM K₂SO₄ considerably stimulated the exudation rate. The application of ABA and IAA (1 mM) also induced a high rate of xylem exudation, K⁺ efflux being simultaneously increased. This revised version was published online in July 2006 with corrections to the Cover Date.     

Apoplastic transport across young maize roots: effect of the exodermis

The uptake of water and of the fluorescent apoplastic dye PTS (trisodium 3-hydroxy-5,8,10-pyrenetrisulfonate) by root systems of young maize (Zea mays L.) seedlings (age: 11–21 d) has been studied with plants which either developed an exodermis (Casparian band in the hypodermis) or were lacking it. Steady-state techniques were used to measure water uptake across excised roots. Either hydrostatic or osmotic pressure gradients were applied to induce water flows. Roots without an exodermis were obtained from plants grown in hydroponic culture. Roots which developed an exodermis were obtained using an aeroponic (=mist) cultivation method. When the osmotic concentration of the medium was varied, the hydraulic conductivity of the root (Lp _r in m³ · m⁻² · MPa⁻¹ · s⁻¹) depended on the osmotic pressure gradient applied between root xylem and medium. Increasing the gradient (i.e. decreasing the osmotic concentration of the medium; range: zero to 40 mM of mannitol), increased the osmotic Lp _r. In the presence of hydrostatic pressure gradients applied by a pressure chamber, root Lp _r was constant over the entire range of pressures (0–0.4 MPa). The presence of an exodermis reduced root Lp _r in hydrostatic experiments by a factor of 3.6. When the osmotic pressure of the medium was low (i.e. in the presence of a strong osmotic gradient between xylem sap and medium), the presence of an exodermis caused the same reduction of root Lp _r in osmotic experiments as in hydrostatic ones. However, when the osmotic concentration of the medium was increased (i.e. the presence of low gradients of osmotic pressure), no marked effect of growth conditions on osmotic root Lp _r was found. Under these conditions, the absolute value of osmotic root Lp _r was lower by factors of 22 (hydroponic culture) and 9.7 (aeroponic culture) than in the corresponding experiments at low osmotic concentration. Apoplastic flow of PTS was low. In hydrostatic experiments, xylem exudate contained only 0.3% of the PTS concentration of the bathing medium. In the presence of osmotic pressure gradients, the apoplastic flow of PTS was further reduced by one order of magnitude. In both types of experiments, the development of an exodermis did not affect PTS flow. In osmotic experiments, the effect of the absolute value of the driving force cannot be explained in terms of a simple dilution effect (Fiscus model). The results indicate that the radial apoplastic flows of water and PTS across the root were affected differently by apoplastic barriers (Casparian bands) in the exodermis. It is concluded that, unlike water, the apoplastic flow of PTS is rate-limited at the endodermis rather than at the exodermis. The use of PTS as a tracer for apoplastic water should be abandoned. Received: 9 October 1997 / Accepted: 5 February 1998     

Water transport in sunflower root systems: effects of ABA, Ca2+ status and HgCl2

Excised 20-d-old sunflower roots (Helianthus annuus L. cv. Sun-Gro 380) with different Ca²⁺ status were used to study the effects of root Ca²⁺ status and abscisic acid (ABA) on the exudation rate (Jv), the hydraulic conductivity of the root (Lpr), the flux of exuded Ca²⁺ (JCa, and the gradient of osmotic pressure between the xylem and the external medium. Jv and Lpr increased in direct proportion to the Ca²⁺ status of the root. Addition of ABA (4 M) at the onset of exudation in the external medium made Jv and Lpr rise, and this effect also increased with the Ca²⁺ status. The effects of HgCl2 and its interaction with ABA on water transport in the root were also studied. Addition of HgCl2 (1 M) 2 h after the onset of exudation in the external medium quickly inhibited Jv, independently of the presence of ABA in the root medium. The results recorded here point to the involvement of ABA and Ca²⁺ in the regulation of root water flow, as well as the existence of aquaporins in the cell membranes of sunflower roots.     

Visualisation of xylem sap flow direction in isolated fine lateral roots and estimation of the xylem sap osmotic potential

Xylem sap outflow from fine lateral roots (FLRs) isolated from hydroponically grown young maize (Zea mays L.) plants was visualized by local brightening of test solutions contrasted with purified Indian ink particles. Flow into the vessels was indicated by the adsorption of Evans Blue in their walls. The fraction of the FLRs able to exude xylem sap in a mineral medium with 30 mM mannitol decreased with increasing incubation time. This change was strongly retarded, when the FLRs were incubated in a medium containing glucose instead of mannitol. There was a broad range of variation of the osmotic potential of the test solutions (Ψ_so), wherein the fraction of the FLRs showing an initially reversed flow of the xylem sap varied between zero and unity. A median (M) of the osmotic potential of the xylem sap in FLRs (Ψ_sx) was estimated. It represents the value of Ψ_so that was lower than Ψ_sx in half of the roots of a sample before their transfer to the test solutions (Ψ_sxo). M was dependent on the osmotic potential of the medium used for growth or pre-incubation of the FLRs. Its value was not dependent on the molecular size of the osmolytes used to adjust Ψ_so, including dextran 8, which is excluded from cell walls. In all of the studied plants, M was lower than the osmotic potential of the xylem sap collected from the root before isolation of the FLRs. To explain this finding it is assumed that FLRs with Ψ_sxo > M had a higher hydraulic conductivity and a larger volume contributed to the exuded sap than those with Ψ_sx < M.     

Analysis of the Exudation Process in Tomato Plants

We started from the supposition that the exudation is broughtabout by two processes: (1) an active transport of salt to thexylem, the ‘salt secretion’, a process that dependson metabolism and is sensitive to inhibitors, and (2) a watertransport effected by the osmotic suction of the sap in thexylem. It was demonstrated that by enhancing the concentrationof the medium the exudation rate rapidly decreases. This isfollowed by a slower rise to a new level which is the consequenceof the salt secretion that continues and enhances the osmoticvalue of the exudation sap. After some time secondary changesof the exudation rate appear. In lowering the concentrationof the medium identical changes occur, but now in the reversedirection. By tracing the influence of addition of substancesdifferent in mol. weight and lipoid solubility it could be establishedthat the gradual change was the result of salt secretion andnot of permeation of the added substance. From this simple suppositionabout the causes of the exudation process we derived formulaefor calculating the salt secretion, the osmotic value of theexudation sap, and the conductivity for water of the root system.Though most observations are in harmony with this concept, thereis a difficulty that too low a value is mostly found for theosmotic value of the exudation sap. The possible causes of thisdeviation are discussed. There is no necessity to accept anactive water secretion as well as active salt secretion sincesalt and water transport cannot be separated. Important factors in exudation are water conductivity of theprotoplasm and salt secretion into the xylem. Both factors areinfluenced by the osmotic value of the outer solution (D.P.D.of the medium) and by the presence of ions in the outer solution.The two factors must have an identical influence on the transportof water into the transpiration stream.     

Evaluation of water stress control with polyethylene glycols by analysis of guttation

The water relations of pepper plants (Capsicum frutescens L.) under conditions conducive to guttation were studied to evaluate the control of plant water stress with polyethylene glycols. The addition of polyethylene glycol 6000 to the nutrient solution resulted in water relations similar to those expected in soil at the same water potentials. Specifically, xylem pressure potential in the root and leaf became more negative during a 24-hour treatment period, while osmotic potential of the root xylem sap remained constant. The decrease in pressure potential was closely correlated with the decrease in osmotic potential of the nutrient solution. In contrast, the addition of polyethylene glycol 400 to the nutrient medium resulted in a reduction of osmotic potential in the root xylem sap; this osmotic adjustment in the xylem was large enough to establish an osmotic gradient for entry of water and cause guttation at a nutrient solution osmotic potential of −4.8 bars. Pressure potential in the root and leaf xylem became negative only at nutrient solution osmotic potentials lower than −4.8 bars. About half of the xylem osmotic adjustment in the presence of polyethylene glycol 400 was caused by increased accumulation of K⁺, Na⁺, Ca²⁺, and Mg²⁺ in the root xylem. These studies indicate that larger polyethylene glycol molecules such as polyethylene glycol 6000 are more useful for simulating soil water stress than smaller molecules such as polyethylene glycol 400.     

Translocation of nitrogen in osmotically stressed wheat seedlings

Wheat (Triticum aestivum L., cv. Drabant) seedlings were grown in a ‘split root’ system where either the whole root system or one root half was subjected to osmotic stress for 24 h, using 200 g polyethylene glycol (PEG, molecular weight 4000) dm^?3 nutrient solution. ¹⁵N-Labelled nitrate was fed to one of the root compartments and total N and ¹⁵N-labelling were measured in plant material and xylem sap. Untreated plants translocated 87% of the N taken up to the shoot, and 10% of this was then retranslocated back to the root. Recalculated on a root nitrogen basis, 36% of the label recovered in the root after 24 h had passed through the shoot. Significant labelling of xylem sap collected from non-labelled roots indicated cycling of organic N through the roots. PEG-treatment of the whole root system caused significant water loss in both roots and shoots. Uptake of nitrate and retranslocation of N to roots were inhibited, whereas cycling of organic nitrogen through the root was still measurable. Treatment of half the root system with PEG had minor effects on shoot water content, but reduced the water content of the treated root part. The total uptake of nitrate by the root system was unaffected, and the effect on the treated root half was comparatively small. Nitrate reductase activity (NRA) declined in PEG-treated roots even if high nitrate uptake rates were maintained. Shoot NRA was unaffected by osmotic stress. The data indicate that the reduction in water content of the root per se has only small effects on nitrate uptake. Major inhibition of nitrate uptake was observed only after treatment of a sufficiently large portion of the root system to given an effect on shoot water content.     

Effect of nitrate on water transfer across roots of nitrogen pre-starved maize seedlings

The addition of 10 mM KNO₃ to the solution bathing the roots of young nitrogen-starved seedlings of Zea mays L. enhanced root water transfer within 15 h, compared with 10 mM KCl addition. The free exudation flux was 2.2–3.9 times higher in excised KNO₃-treated roots than in KCl-treated ones. Cryo-osmometry data for xylem sap suggested that, compared with chloride, nitrate treatment increased the steady solute flux into the xylem, but did not modify the osmotic concentration of sap. Root growth was not significantly modified by nitrate within 15 h. Root hydraulic conductances were measured by using either hydrostatic-pressure or osmotic-gradient methods. During hydrostatic experiments, the conductance (k_p), which is thought to refer mainly to the apoplasmic pathway, was 1.6 times larger in KNO₃-than in KCl-treated plants. From experiments in which polyethylene glycol (PEG) 8000 was used as external osmolyte, osmotic conductances (k_s) were found to be smaller by 5–20 times than k_p for the two kinds of plants. The KCl-treated roots were characterized by a low k_s which was the same for influx or efflux of water. By contrast, KNO₃-treated roots exhibited two distinct conductances k_s1 and k_s2, indicating that influx of water was easier than efflux when the water flow was driven by the osmotic pressure gradient. Infiltration of roots with KNO₃ solution supported the idea that nitrate might enhance the efficiency of the cell-to-cell pathway. The low k_s value of KCl-treated roots and the existence of two contrasting k_s values (k_s1 and k_s2) for KNO₃-treated roots are discussed in terms of reversible closing of water channels.     

Water relations in silver birch during springtime: How is sap pressurised?

• Positive sap pressures are produced in the xylem of birch trees in boreal conditions during the time between the thawing of the soil and bud break. During this period, xylem embolisms accumulated during wintertime are refilled with water. The mechanism for xylem sap pressurization and its environmental drivers are not well known.
• We measured xylem sap flow, xylem sap pressure, xylem sap osmotic concentration, xylem and whole stem diameter changes, and stem and root non‐structural carbohydrate concentrations, along with meteorological conditions at two sites in Finland during and after the sap pressurisation period.
• The diurnal dynamics of xylem sap pressure and sap flow during the sap pressurisation period varied, but were more often opposite to the diurnal pattern after bud burst, i.e. sap pressure increased and sap flow rate mostly decreased when temperature increased. Net conversion of soluble sugars to starch in the stem and roots occurred during the sap pressurisation period. Xylem sap osmotic pressure was small in comparison to total sap pressure, and it did not follow changes in environmental conditions or tree water relations.
• Based on these findings, we suggest that xylem sap pressurisation and embolism refilling occur gradually over a few weeks through water transfer from parenchyma cells to xylem vessels during daytime, and then the parenchyma are refilled mostly during nighttime by water uptake from soil. Possible drivers for water transfer from parenchyma cells to vessels are discussed. Also the functioning of thermal dissipation probes in conditions of changing stem water content is discussed.

     

Quantification of the Daily Cytokinin Transport from the Root to the Shoot of Urtica dioica L.*

Cytokinins are predominantly root-born phytohormones which are distributed in the shoot via the xylem stream. In the hormone message concept they are considered as root signals mediating the transport of the photosynthates to the various sinks of a plant. In this paper the cytokinin relations of Urtica dioica L., the stinging nettle, are described, based on the daily flux from the roots to the shoot. Trans-zeatin-type cytokinins predominate in the various tissues of Urtica (Wagner and Beck, 1993), and accordingly trans-zeatin riboside and trans-zeatin are the forms transported by the xylem sap. The daily time-course of cytokinin concentration in root pressure exudates and in xylem sap collected from a petiole after pressurizing the root bed showed high concentrations in the morning, followed by a substantial drop to a level of 15–30% of the initial concentration which was then maintained during the afternoon. This time-course is interpreted as resulting from continuous synthesis and exudation of cytokinins into the xylem fluid of the roots whose cytokinin concentration is then modified by the dynamics of the transpiration stream. Loading of cytokinins into the xylem sap could be enhanced several times by increasing the flux rate of the xylem stream to the maximal transpiration rate when a maximum export rate was reached. The total daily cytokinin gain by the shoot depended on the nitrogen status of the plant. Roots of Urtica plants grown on a sufficient nitrogen supply had a significantly higher cytokinin content and exuded more cytokinins into the shoot than those of plants raised under nitrogen shortage. A positive correlation was found between the steady rates of cytokinin export measured during the afternoon and the shoot to root-ratios of biomass which, in turn, corresponded to the nitrogen status of the plants.     

Cytoplasmic streaming in the root cortex and its role in the delivery of potassium to the shoot

The influence of cytochalasin B (CB), a potent inhibitor of cytoplasmic streaming, on ⁸⁶Rb-labelled K⁺ translocation by detopped Lycopersicon esculentum Mill., Cucumis sativus L. and Zea mays L. plants was examined by measuring the radioactivity in xylem exudate before and after the addition of CB to the medium bathing the roots. CB caused complete cessation of cytoplasmic streaming in root segments within 15 min but was without effect on either total ⁸⁶Rb uptake or exudation. Thus factors other than cytoplasmic streaming limit the movement of K⁺ across the symplast of the root of higher plants.     

Perfusion of onion root xylem vessels: a method and some evidence of control of the pH of the xylem sap

We describe a method for perfusing the xylem in the stele of excised onion roots with solutions of known composition under a pressure gradient. Tracer studies using [¹⁴C] polyethylene glycol 4000 and the fluorescent dye, Tinopal CBSX, indicated that perfusing solutions passed exclusively through the xylem vessels. The conductance of the xylem was small over the apical 100 mm of the root axis but increased markedly between 100 and 200 mm. Unbuffered perfusion solutions supplied in the range pH 3.7–7.8 emerged after passage through the xylem adjusted to pH 5.2–6.0, indicating the presence of mechanisms for absorbing or releasing protons. This adjustment continued over many hours with net proton fluxes apparently determined by the disparity between the pH of the perfusion solution and the usual xylem sap pH of about 5.5. Mild acidification of the xylem sap by buffered perfusion solutions increased the release of ⁸⁶Rb (K⁺) and ³⁵SO₄ ^2- from the stelar tissue into the xylem stream. The ion-transporting properties of onion roots seemed little changed by excision from the bulbs, or by removal of the apical zones of the root axis. The pH of sap produced by root pressure resembles that found in the outflow solutions of perfused root segments.     

Form of aluminium for uptake and translocation in buckwheat (Fagopyrum esculentum Moench)

 The forms of Al for uptake by the roots and translocation from the root to the shoot were investigated in a buckwheat (Fagopyrum esculentum Moench, cv. Jianxi) that accumulates Al in its leaves. The Al concentration in the xylem sap was 15-fold higher in the plants exposed to AlCl₃ than in those exposed to an Al-oxalate (1:3) complex, suggesting that the roots take up Al in the ionic form. The Al concentration in the xylem sap was 4-fold higher than that in the external solution after a 1-h exposure to AlCl₃ solution and 10-fold higher after a 2-h exposure. The Al concentration in the xylem sap increased with increasing Al concentration in the external solution. The Al uptake was not affected by a respiratory inhibitor, hydroxylamine, but significantly inhibited by the addition of La. These results suggest that Al uptake by the root is a passive process, and La³⁺ competes for the binding sites for Al³⁺ on the plasma membrane. The form of Al in the xylem sap was identified by ²⁷Al-nuclear magnetic resonance analysis. The chemical shift of ²⁷Al in the xylem sap was around 10.9 ppm, which is consistent with that of the Al-citrate complex. Furthermore, the dominant organic acid in the xylem sap was citric acid, indicating that Al was translocated in the form of Al-citrate complex. Because Al is present as Al-oxalate (1:3) in the root, the present data show that ligand exchange from oxalate to citrate occurs before Al is released to xylem. Received: 10 December 1999 / Accepted: 3 February 2000     

Na~ and Water Uptake in Relation to the Radial Reflection Coefficient of Root in Arrowleaf Saltbush Under Salt Stress

The response of halophyte arrowleaf saltbush(Atriplex triangularis Willd)plants to a gradient of salt stress were investigatedwith hydroponically cultured seedlings.Under salt stress,both the Na~ uptake into root xylem and negative pressures inxylem vessels increased with the elevation of salinity(up to 500 mol/m~3)in the root environment.However,the increment innegative pressures in root xylem far from matches the decrease in the osmotic potential of the root bathing solutions,evenwhen the osmotic potential of xylem sap is taken into consideration.The total water potential of xylem sap in arrowleafsaltbush roots was close to the osmotic potential of root bathing solutions when the salt stress was low,but a progressivelyincreased gap between the water potential of xylem sap and the osmotic potential of root bathing solutions was observedwhen the salinity in the root environment was enhanced.The maximum gap was 1.4 MPa at a salinity level of 500 mol/m~3without apparent dehydration of the tested plants.This discrepancy could not be explained with the current theories inplant physiology.The radial reflection coefficient of root in arrowleaf saltbush decreased with the enhanced salt stress wasand accompanied by an increase in the Na~ uptake into xylem sap.However,the relative Na~ in xylem exudates based onthe corresponding NaCl concentration in the root bathing solutions showed a tendency of decrease.The results showedthat the reduction in the radial reflection coefficient of roots in the arrowleaf saltbush did not lead to a mass influx of NaClinto xylem when the radial reflection coefficient of the root was considerably small;and that arrowleaf saltbush could usesmall xylem pressures to counterbalance the salt stresses,either with the uptake of large amounts of salt,or with thedevelopment of xylem pressures dangerously negative.This strategy could be one of the mechanisms behind the highresistance of arrowleaf saltbush plants to salt stress.     

Stomatal Behaviours of Aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis>) Plants in Response to Low Root Temperature in Hydroponics

Hydroponic-grown seedlings of aspen (Populus tremuloides Michx.) were used to investigate how low root temperatures (5°C) affect stomatal conductance and water relations. An isohydric manner of the stomatal behaviour was found with the seedlings when their roots were subjected to the low temperature. Stomatal conductance rapidly and dramatically reduced in response to the low root temperature, while the xylem water potential did not significantly alter. Under the low root temperature, pH value of the xylem sap increased from 6.15 to 6.72 within the initial 4 h, while abscisic acid (ABA) concentration increased by the eighth hour of treatment. K⁺ concentration of the xylem sap significantly decreased within the 8th h and then reversed by the 24th h. The ion change was accompanied by a decrease and then an increase in the electrical conductivity, and an increase and then a decrease in the osmotic potential. The tempo of physiological responses to the low root temperature suggests that the rapid pH change of the xylem sap was the initial factor which triggered stomatal closure in low temperature-treated seedlings, and that the role of the more slowly accumulating ABA was likely to reinforce the stomatal closure. Xylem sap from the seedlings subjected low root temperature affected stomatal aperture on leaf discs when they were floated on the sap solution. The stomatal aperture correlated (P = 0.006) with the changed pattern of [K⁺] in the sap while the range of pH or ABA found in the xylem sap did not influence stomatal aperture of leaf discs in solution. The effect of xylem sap on stomatal aperture on leaf discs was different from on stomatal conductance in the intact seedlings. Comparison was made with previous study with the soil-grown seedlings.     

Growth of Zea mays L. plants with their seminal roots only. Effects on plant development, xylem transport, mineral nutrition and the flow and distribution of abscisic acid (ABA) as a possible shoot to root signal

Maize (Zea mays L.) was grown in quartz sand culture eitherwith a normal root system (controls) or with seminal roots only(‘single-rooted’). Development of adventitious rootswas prevented by using plants with an etiolated mesocotyl andthe stem base was positioned 5–8 cm above the sand. Eventhough the roots of the single-rooted plants were sufficientlysupplied with water and nutrients, the leaves experienced waterdeficits and showed decreased transpiration as trans plrationalwater flow was restricted by the constant number of xylem vesselspresent in the mesocotyl. As a consequence of this restriction,transpirational water flow velocities in the metaxylem vesselsreached mean values of 270 m h^–1 and phloem transportvelocities of 5.2 m h^–1. Despite limited xylem transportmineral nutrient concentrations in leaf tissues were not decreasedin single-rooted plants, but shoot and particularly stem developmentwas somewhat inhibited. Due to the lack of adventitious rootsthe shoot:root ratio was strongly increased in the single-rootedplants, but the seminal roots showed compensatory growth comparedto those in control plants. Consistent with decreased leaf conductance,ABA concentrations in leaves of single-rooted plants were elevatedup to 10-fold, but xylem sap ABA concentrations in these plantswere lower than in controls, in good agreement with the well-wateredconditions experienced by the seminal roots. Surprisingly, however,ABA concentrations in tissues of the seminal roots of the single-rooted plants were clearly increased compared to the controls,presumably due to increased ABA import via phloem from the water-stressedleaves. The results are discussed in relation to the role ofABA as a shoot to root signal. Key words: Zea mays, seminal roots, plant development, xylem transport, mineral nutrition, ABA, shoot-to-root signal     

Na^＋ and Water Uptake in Relation to the Radial Reflection Coefficient of Root in Arrowleaf Saltbush Under Salt Stress

The response of halophyte arrowleaf saltbush (Atriplex triangularis Willd) plants to a gradient of salt stress were investigated with hydroponically cultured seedlings. Under salt stress, both the Na+ uptake into root xylem and negative pressures in xylem vessels increased with the elevation of salinity (up to 500 mol/m3) in the root environment. However, the increment in negative pressures in root xylem far from matches the decrease in the osmotic potential of the root bathing solutions, even when the osmotic potential of xylem sap is taken into consideration. The total water potential of xylem sap in arrowleaf saltbush roots was close to the osmotic potential of root bathing solutions when the salt stress was low, but a progressively increased gap between the water potential of xylem sap and the osmotic potential of root bathing solutions was observed when the salinity in the root environment was enhanced. The maximum gap was 1.4 MPa at a salinity level of 500 mol/m3 without apparent dehydration of the tested plants. This discrepancy could not be explained with the current theories in plant physiology. The radial reflection coefficient of root in arrowleaf saltbush decreased with the enhanced salt stress was and accompanied by an increase in the Na+ uptake into xylem sap. However, the relative Na+ in xylem exudates based on the corresponding NaCl concentration in the root bathing solutions showed a tendency of decrease. The results showed that the reduction in the radial reflection coefficient of roots in the arrowleaf saltbush did not lead to a mass influx of NaCl into xylem when the radial reflection coefficient of the root was considerably small; and that arrowleaf saltbush could use small xylem pressures to counterbalance the salt stresses, either with the uptake of large amounts of salt, or with the development of xylem pressures dangerously negative. This strategy could be one of the mechanisms behind the high resistance of arrowleaf saltbush plants to salt stress.     

Is xanthine dehydrogenase involved in response of pea plants (<Emphasis Type="Italic">Pisum sativum</Emphasis> L.) to salinity or ammonium treatment?

The effect of salinity and different nitrogen sources on the level of xanthine dehydrogenase (XDH) activity in roots and leaves of pea plants was investigated. Two bands of xanthine dehydrogenase activity (XDH-R2, XDH-R3) were detected in roots after native PAGE and staining with hypoxanthine as substrate. Only one band of XDH activity (XDH-L1) was detected in leaf extracts. Within leaves of three different ages the highest XDH activity was detected in young leaves both under control as well as stress conditions. Salinity did not affect significantly the activity of XDH in pea roots, however, depressed XDH activity in leaves. A significant increase of XDH activity both in roots and leaves was observed only when ammonium was applied as the sole N source. Increased concentration of ureides in the xylem sap of pea plants was observed for both ammonium and high salt treatments, although the higher content of ureides in the xylem sap of 100 mM NaCl treated plants may be rather a result of lower rate of exudation from roots than of increased root ureide biosynthesis. Thus, the changes of root and leaf XDH activity in pea plants seem to be tightly correlated with ureide synthesis that is induced by NH ₄ ⁺ , the product of N fixation, and rather than by salinity. A contribution of pea XDH in increased oxygen species or uric acid production under saline conditions seems to be less than likely.