Regulation of K+ Influx in Barley: Evidence for a Direct Control of Influx by K+ Concentration of Root Cells

Siddiqi, M. Y. and Glass, A. D. M. 1987. Regulation of K⁺ influxin barley: Evidence for a direct control of influx by K⁺ concentrationof root cells.—J. exp. Bot. 38: 935–947. The kinetics of K⁺ (⁸⁶Rb⁺) influx into intact roots of barley(Hordeum vulgare L. cv. Fergus) seedlings having different combinationsof root and shoot [K⁺], different growth rates and differentroot:shoot weight ratios were studied. K⁺ influx was stronglycorrelated with root [K⁺]; shoot [K⁺], growth rates, and root:shoot ratios appeared to have little effect on K⁺ influx. Adetailed study showed that both V_max and K_m for K⁺ influx wereaffected by root [K⁺] but not by shoot [K⁺]. We have suggestedthat factors such as growth rates and root: shoot ratio mayaffect K⁺ influx indirectly primarily via their influence onroot factors such as root [K⁺]. We have reiterated that othertypes of kinetic control, e.g. increased or decreased synthesisof ‘carrier systems’, may operate in addition todirect (allosteric?) control of K⁺ influx by root [K⁺]. Thenegative feedback signal from root [K⁺] appeared to be the primeeffector in the regulation of K⁺ influx. Key words: Barley, K⁺ influx     

Ionic Relations of Garden Orache, A triplex hortensis L.: Growth and Ion Distribution at Moderate Salinity and the Function of Bladder Hairs

The growth of garden orache, A triplex hortensis was studiedunder conditions of mild NaCl or Na₂SO₄ salinity. Growth, drymatter production and leaf size were substantially stimulatedat 10 mM and 50 mM Na⁺ salts. Increased growth, however, appearedto be due to a K⁺-sparing effect of Na⁺ rather than to salinityper se. The distribution of K⁺ and Na⁺ in the plant revealeda remarkable preference for K+ in the roots and the hypocotyl.In the shoot the K/Na ratio decreased strongly with leaf age.However, the inverse changes in K⁺ and Na⁺ content with leafage were dependent on the presence of bladder hairs, which removedalmost all of the Na⁺ from the young leaf lamina. Measurementsof net fluxes of K⁺ and Na⁺ into roots and shoots of growingAtriplex plants showed a higher K/Na selectivity of the netion flux to the root compared to the shoot. With increasingsalinity the selectivity ratio S_{K, Na}^* of net ion fluxes tothe roots and to the shoots was increased. The data suggestthat recirculation of K⁺ from leaves to roots is an importantlink in establishing the K/Na selectivity in A. hortensis plants.The importance of K⁺ recirculation and phloem transport forsalt tolerance is discussed. Key words: Atriplex hortensis, Salinity, Potassium, Sodium, K⁺ retranslocation, Bladder hairs, Growth stimulation     

Diurnal Rb+ Transport from Roots to the Laminar Pulvinus in Phaseolus vulgaris L.

The primary leaves of kidney bean (Phaseolus vulgaris L.) openunder light and close in the dark by the deformation of thepulvinus resulting from diurnal distribution changes of K⁺,Cl^–, organic acid (or H⁺) and NO₃^–. When Rb⁺ was added as a tracer of K⁺ to the seedlings throughtheir roots, it was transported to the pulvinus cells duringthe light period but not during the dark period. Transpirationoccurred vigorously in the light but almost stopped in the dark.We concluded that Rb⁺ absorbed by the roots was carried to thepulvinus by the transpiration stream. Phaseolus vulgaris L., pulvinus, Rb⁺, diurnal transport transpiration stream     

Response of Wheat Seedlings to Low O2 Concentrations in Nutrient Solution: II. K +/Na+ SELECTIVITY OF ROOT TISSUES9

This paper reports the effects of low O₂ concentration (0–01,0–055, and 0.115mol m^–3) in nutrient solutions onK⁺/Na⁺ selectivity of growing and mature root tissues of 6-to 8-d-old, intact, wheat (Triticum aestivum cv. Gamenya) seedlings. Increases in anaerobic catabolism and decreases in O₂ uptake,K⁺ uptake and K⁺/Na⁺ selectivity were all more pronounced and/oroccurred at higher external O₂ concentrations in the apex (0–2mm) than in the expanding tissues (2–4 mm); these growingtissues were, in turn, more affected than the expanded tissuesof the roots (4–12 mm). Selectivity for K⁺ over Na⁺ in roots and shoots was particularlysensitive to O₂ deficiency. For example, in apical tissues (0–2mm) K ⁺ /Na⁺ selectivity was already reduced at 0.115 mol m^–3O₂, yet at this O₂ concentration there was no effect on eithergrowth or (K⁺/Na⁺) uptake. Upon transfer from 0.01 to 0.26 mol m^–3 O₂, a detailedstudy of the 12 mm root tips showed that 70% of these tips regainedhigh (K⁺ + Na⁺) concentrations and K⁺/^Na+ ratios. In contrast,there was no recovery in the remaining 30% of the 12 mm roottips. Net K⁺ transport to the shoots during the period afterre-aeration was negative for the population as a whole. Theseverity of these effects supports the view that the root tipsand the stele were more susceptible to O2 deficiency than wasthe cortex of the fully-developed root tissues. Key words: Hypoxia, K⁺/Na⁺ selectivity, expanded and expanding tissues     

Application of 39K NMR Spectroscopy to Potassium Transport in Mung Bean Root Tips

The intracellular K⁺ concentration and its change in mung bean[Vigna mungo (L.) Hepper] root tips were investigated non-invasivelywith ³⁹K nuclear magnetic resonance spectroscopy using a membraneimpermeable shift reagent, dysprosium (III) tripolyphosphate[Dy(PPP_i)^7–₂]. The K⁺ resonance was shifted to highermagnetic field in proportion to the concentration of the shiftreagent. In addition to a reference capillary peak for measuringthe K⁺ concentration, two well-resolved peaks (intra- and extracellularK⁺ resonances) were observed in the 39K NMR spectra of mungbean root tips. The intracellular K⁺ concentration was determinedto be 41 mM, which was similar to the value obtained by flamephotometry. When 20 mM KCl was added to the external medium,the intensity of the intracellular K⁺ resonance gradually increasedand the net K⁺ uptake rate was calculated to be 4.1 micromolesper gram fresh weight per hour. After removal of KCl from theperfusion medium, the intracellular K⁺ concentration considerablydecreased. With ³¹P NMR method, 2.5 mM Dy(PPP_j)^7–1₂ and20 mM KCl had little effect on the ATP level in the cells. Wehave indicated that the ³⁹K NMR method can be used to determinethe K⁺ levels and net fluxes of the K⁺ transport in perfusedroot tips successively. (Received April 6, 1988; Accepted September 29, 1988)     

External Potassium Supply is not Required for Root Growth in Saline Conditions: Experiments with Ricinus communis L. Grown in a Reciprocal Split-Root System

Ricinus communis L. (castor bean) plants were grown in the absence(control) and in the presence of 100molm^–3NaCl with areciprocal split-root system, in which K⁺ was supplied to oneand NO₃^– to the other part of the root system. In theseplants shoot and, to a lesser extent, total root growth wereinhibited compared to plants with non-split roots. Without andwith NaCl, growth of roots receiving NO₃^– but noK⁺ (‘minusK/plus N-roots’) was substantially more vigorous thanunder the reverse conditions (‘plus K/minus N-roots¹).100mol m^–3 NaCl inhibited growth of minus K/plus N-roots¹to the same extent as that of non-split roots, indicating thatexternally supplied K⁺ was not required for root growth undersaline conditions. In growth media without added K⁺ the rootdepleted the external low K ⁺ levels resulting from chemicalsdown to a minimum value C_mln (1.0 to 1.4 mmol m^–3); inthe presence of 100 mol m^–3 NaCl, C_min, was higher (10–18mmol m^–3) and resulted from an initial net loss of K ⁺.C_min, was pH-dependent The distribution of K⁺, Na⁺ and Mg²⁺along the root was measured. In meristematic root tissues, K⁺ concentrations were scarcely affected by external K⁺ or byNaCl, where Na ⁺ concentrations were low, but somewhat elevatedat low external K+ and/or high NaCl. In differentiated, vacuolatedtissues K ⁺ concentrations were low and Na⁺ concentrations high,if K ⁺ was not supplied externally and/or NaCl was present.The longitudinal distribution of ions within the root was usedto estimate cytoplasmic and vacuolar ion concentrations. Thesedata showed a narrow homoeostasis of cytoplasmic K+ concentrations(100–140 mol m^–3) independent of external K ⁺ supplyeven in the presence of 100 mol m ^–3 NaCl. CytoplasmicNa ⁺ concentrations were maintained at remarkably low levels.Hence, external K⁺ concentrations above C_min, were not requiredfor maintaining K/Na selectivity, i.e. for controlling Na⁺ entry.The results are discussed with regard to mechanisms of K/Naselectivity and to the importance of phloem import of K⁺ forsalt tolerance of roots and for cytoplasmic K⁺ homoeostasis. Key words: Ricinus communis, nitrate, potassium, root (split-root), salt tolerance, phloem transport     

Effect of K+, its Counter Anion, and pH on Sodium Efflux from Barley Root Tips

Sodium efflux from ²²Na⁺-loaded root tips root tips of Hordeumvulgare L. was markedly increased by replacing 10mM Na₂SO₄ inthe washing solution by K₂SO₄ with the same electrical conductivity.This increase was inhibited by both an uncoupler and an inhibitorof oxidative phosphorylation but not by ouabain. Potassium ionsdid not enhance Na⁺ efflux in the presence of a rapidly absorbedcounter anion, such as Cl^–, instead of . Efflux of ²²Na⁺ could also be enhanced by a low pH in theabsence of K⁺; this was prevented by uncouplers, but not byan inhibitor of the mitochondrial ATPase. It seems that K⁺ indirectly enhances Na⁺ efflux. It is suggestedthat metabolic K⁺ uptake in excess of the counter anion resultsin a proton gradient across the plasmalemma (acid outside) inducingH⁺/Na⁺ antiport.     

Effects of Growth and Assay Temperatures on Unidirectional K+ Fluxes in Roots of Rye (Secale cereale)

The effects of growth and assay temperature on unidirectionalK⁺ fluxes in excised roots of rye (Secale cereale cv. Rheidol)were studied using ⁸⁶Rb⁺ as a tracer. Both K⁺ influx to thevacuole, estimated as K⁺ uptake between 3 and 12 h after transferof unlabelled roots to radioactive solution, and movement ofK⁺ to the xylem were determined directly. Other fluxes weredetermined on excised roots of plants, which had been labelledwith ⁸⁶Rb⁺ since germination, by conventional triple exponentialefflux analysis. When assayed at 20°C, roots of plants previously grown at20°C(WG roots) had lower rates of net K⁺ uptake than rootsof low temperature-acclimated plants, grown with a temperaturediferential between roots (87°C) and shoots (20°C) eithersince germination (DG roots) or for 3 d prior to experiments(DT roots). This resulted from a greater unidirectional K⁺ effluxacross the plasma membrane and a reduced K⁺ flux to the xylemin WG roots, compared to DG or DT roots, rather than a decreasein unidirectional K⁺ influx or a decrease in the net K⁺ fluxto the vacuole. Indeed, although WG roots had lower rates ofK⁺ influx and K⁺ efflux across the tonoplast at 20°C thanDG or DT roots, roots of plants from all growth temperaturetreatments showed an equivalent net K⁺ flux to the vacuole. Although all unidirectional K⁺ fluxes in roots from plants grownunder all temperature regimes were reduced by lowering the temperatureof the root, these fluxes were differentially affected in rootsof plants from contrasting growth temperature treatments. Rapidcooling to 8°C of WG roots resulted in a lower rate of K+influx and a transient increase in K⁺ efflux across both theplasma membrane and tonoplast, compared to DG and DT roots.Furthermore, since the K⁺ flux to the xylem was lower in WGroots, the net K⁺ uptake at 8°C into WG roots was considerablyreduced compared to DG and DT roots. These results suggest thatlow temperature-acclimation of K⁺ fluxes in rye roots may involvea reduction in the temperature sensitivity of K⁺ influx anda curtailment of K⁺ efflux across both the plasma membrane andtonoplast at low temperatures. Key words: K⁺influx, K⁺ efflux, low temperature, potassium, rye (Secale cereale cv. Rheidol)     

Hypoxia Induces Membrane Depolarization and Potassium Loss from Wheat Roots but does not Increase their Permeability to Sorbitol

This paper deals with the responses of roots of wheat {Triticumaestivum L.) to hypoxia with special emphasis on the effectsof severe O2 deficiency on membrane integrity, loss of K+ fromthe root and root membrane potentials. Seminal and crown roots of 26-d-old plants exposed to severehypoxia (0.003 mol O₂ m^–3) for 3 h or 10 d prior to excisionand subsequently exposed to hypoxic solutions, had slightlylower rates of sorbitol influx and a slightly smaller apparentfree space than roots in aerated solutions. These results indicatethat neither a few hours nor a 10-d exposure to hypoxia hadadverse effects on the membrane integrity of the bulk of thecells in the roots. However, both 6-d-old seedlings and 26-d-oldplants lost K⁺ from the roots following their transfer fromaerated to hypoxic nutrient solutions. In the 26-d-old plants,which were of high nutritional status, there was a net K⁺ effluxfrom the roots to the external solution. In contrast, with the6-d-old seedlings, which were of low nutritional status, thedecrease in the K⁺ content of the roots was smaller than thenet K⁺ uptake to the shoots. Exposure of excised roots to 0.008 mol O₂ m^–3caused arapid and reversible membrane depolarization from –120to ––80 mV. These data and the magnitude of thenet effluxes strongly suggest that K⁺ losses during the earlystages of hypoxia are due to membrane depolarization ratherthan to increases in the permeability of membranes to K ⁺. Key words: Hypoxia, membrane integrity, membrane potentials, seminal and crown roots     

The Influence of NO-3 and NH+4 Nutrition on the Gas Exchange Characteristics of the Roots of Wheat (Triticum aestivum) and Maize (Zea mays) Plants

Respiratory oxygen consumption by roots was 1·4- and1·6-fold larger in NH⁺₄-fed than in NO^-₃-fed wheat (Triticumaestivum L.) and maize (Zea mays L.) plants respectively. Higherroot oxygen consumption in NH⁺₄-fed plants than in NO^-₃-fedplants was associated with higher total nitrogen contents inNH⁺4-fed plants. Root oxygen consumption was, however, not correlatedwith growth rates or shoot:root ratios. Carbon dioxide releasewas 1·4- and 1·2-fold larger in NO⁺3-fed thanin NH⁺₄-fed wheat and maize plants respectively. Differencesin oxygen and carbon dioxide gas exchange rates resulted inthe gas exchange quotients of NH^-₄-fed plants (wheat, 0·5;maize, 0·6) being greatly reduced compared with thoseof NO^-₃-fed plants (wheat, 1·0; maize, 1·1). Measuredrates of HCO^-₃ assimilation by PEPc in roots were considerablylarger in 4 mM NH⁺₄-fed than in 4 NO^-₃ plants (wheat, 2·6-fold;maize, 8·3-fold). These differences were, however, insufficientto account for the observed differences in root carbon dioxideflux and it is probable that HCO^-₃ uptake is also importantin determining carbon dioxide fluxes. Thus reduced root extension in NH⁺₄-fed compared with NO^-₃-fedwheat plants could not be ascribed to differences in carbondioxide losses from roots.Copyright 1993, 1999 Academic Press Triticum aestivum, wheat, Zea mays, maize assimilation, ammonium assimilation, root respiration     

Oxidative Damages and Repair in Euglena gracilis Exposed to Ozone II. Membrane Permeability and Uptake of Metabolites

Significant injuries to the plasma membrane were detected inEuglena gracilis cells during ozone exposure (240 µ1.liter¹,delivery rate of l µmol.min^–1), as assessed by measuringthe alterations of vitamin B₁₂ and acetate uptakes and the leakageof intracellular K⁺ (Rb⁺). A rapid decrease in the uptake ofvitamin B₁₂ and acetate was observed within 15 min of treatment,indicating that both transport systems are very sensitive toO₃. On the other hand, the leakage of intracellular K⁺ ions,as measured by the efflux of ⁸⁶Rb⁺ from prelabelled cells, couldonly be detected after 30 min of O₃ exposure. These resultssuggest that the initial metabolic symptoms of injury is atthe level of the two transport systems examined and that thealteration of the membrane permeability to K⁺ ions appears asa second step in the cascade of oxidative events at the plasmamembrane level. When Euglena cells were allowed to recover underautotrophic growth conditions following O₃ treatment, vitaminB₁₂ and ⁸⁶Rb⁺ (K⁺) ions uptakes returned gradually to controllevel within 5 h of the recovery period. Acetate uptake returnedto control level at a slower rate and needed 20 h for completerecovery. These results indicate that the cells were able toactively repair most of the initial oxidative damages inducedby O₃. The metabolic significance of the repair mechanism(s)is discussed. (Received December 25, 1989; Accepted July 23, 1990)     

The influence of salinity on the utilization of root anaplerotic carbon and nitrogen metabolism in tomato seedlings

In hydroponically grown Lycopersicon esculentum (L.) Mill. cv.F144 the site of NO₃^– reduction and assimilation withinthe plant was shifted from the shoot to the root by salinity.Uptake of NO₃^– from the root solution was strongly inhibitedby salinization. Consequently, NO₃^– concentrations inthe leaf, stem and root tissues as well as the nitrate reductaseactivities of the leaves were lower in salinized than in controlplants. Lower NO₃^–, but higher reduced-N, concentrationswere observed in the xylem sap as a result of the enhanced participationof the root in NO₃^– reduction in salinized plants. Lowerstem K⁺ concentrations and leaf malate concentrations were foundin salinized compared to control plants which indicates reducedfunctioning of the K⁺–shuttle in the salinized plants. Incorporation of inorganic carbon by the root was determinedby supplying a pulse of NaH¹⁴CO₃ followed by extraction andseparation of the labelled products on ion exchange resins.The rate of H¹⁴CO₃^– incorporation was c. 2-fold higherin control than in salinized plants. In salinized plants theproducts of H¹⁴CO₃^– incorporation within the roots werediverted into amino acids, while the control plants divertedrelatively more ¹⁴C into organic acids. Products of inorganiccarbon incorporation in the roots of salinized plants providean anaplerotic source of carbon for assimilation of reducedNO₃^– into amino acids, while in control plants the productswere predominantly organic acids as part of mechanisms to maintainionic balance in the cells and in the xylem sap. Key words: Tomato, nitrate, PEPc, respiration, salinity     

The regulation of K+ influx into roots of rye (Secale cereale L.) seedlings by negative feedback via the K+ flux from shoot to root in the phloem

The uptake of K⁺ by plant roots is matched to the demand forK⁺ for growth. The growing shoot must communicate its K⁺ requirementto the root. It has been suggested that this might be effectedby varying the amount of K⁺ retranslocated from the shoot tothe root via the phloem. It is predicted that less K⁺ is returnedto the roots in K⁺-deficient plants and that this promotes compensatoryK⁺ uptake from the external medium. These experiments addressthis hypothesis. Rye (Secale cereale L.) was grown hydroponically in completenutrient solutions containing either 100 aM or 400 µMK⁺. Plant development, shoot fresh weight (FW) and plant drymatter accumulation did not differ between seedlings grown atthese K⁺ concentrations. However, root FW was lower in seedlingsgrown in solutions containing 100 µM K⁺, which resultedin a greater shoot/root FW ratio. Seedlings from both treatmentshad similar shoot K⁺ concentrations, but the root K⁺ concentrationof seedlings grown In solutions containing 100 µM K⁺ wasless than their counterparts grown at 400 µM K⁺. When assayed at the same K⁺ concentration, unidirectional K⁺(⁸⁶Rb⁺) influx into 14-d-old seedlings grown with 100 µMK⁺ in the nutrient solution was greater than that into seedlingsgrown with 400 µM K⁺ in the nutrient solution, indicatingan increased K⁺ influx capacity in the former. Furthermore,K⁺ (⁸⁶Rb⁺) influx into seedlings grown and assayed at 100 µMK⁺ was greater than that into seedlings grown and assayed at400 µM K⁺. Since net K⁺ uptake was lower in the seedlingsgrown at 100 µM K⁺, this indicates a greater unidirectionalK⁺ efflux from roots in solutions containing 100 µM K⁺. An empirical model, based on the immobility of calcium in thephloem, was used to describe quantitatively K⁺ fluxes in seedlings14 d after sowing. As primary data, the composition of xylemsap and the accumulation of elements in root and shoot tissueswere determined. Xylem sap was collected either as root-pressureexudate or from excised roots immersed in nutrient solutionand subjected to a pneumatic pressure of 0.4 MPa. The K:Ca ratioin these saps differed, and led to contrasting conclusions concerningthe effect of K⁺ nutrition on the recirculation of K⁺. Basedon the K:Ca ratio in the sap obtained following the applicationof pneumatic pressure, which is thought to resemble that ofintact transpiring plants, it was calculated that the K⁺ fluxfrom the shoot to the root was higher in seedlings maintainedin solutions containing higher K⁺ concentrations. This suggeststhat a negative feedback mechanism based on K⁺ recirculationfrom the shoot to the root via the phloem could be a primarysignal decreasing K⁺ influx. Key words: K⁺ influx, K⁺ recirculation, regulation, root, rye, Secale cereale L     

Radial Movement of Cations across Aerenchymatous Roots of Zea mays Measured by Electron Probe X-Ray Microanalysis

The radial movements of Rb⁺ and Sr²⁺ as analogues for K⁺ andCa²⁺ were followed by electron probe X-ray microanalysis inaerenchymatous roots of maize (Zea mays L. cv. LG 11). The objectwas to determine the possible pathways by which ions can reachthe xylem when degeneration and lysis of much of the root cortexhas taken place during formation of numerous gas-filled spacesin aerenchymatous roots. After 1, 6 and 24 h uptake of Rb⁺ andSr²⁺ from a full strength nutrient solution containing K⁺ andCa²⁺, the distribution of these elements was examined. Transverseroot sections, prepared by cryostat sectioning and lyophilization,were used for electron probe X-ray microanalysis. If the cellwalls lining gas-filled spaces become suberized (as reportedin earlier literature), this development might be expected toretard ion movement, but we found that Rb⁺ or Sr²⁺ could migrateas readily in the radial wall residues remaining after corticalcell lysis, as in the walls of intact cortical cells. The distributionsof Rb⁺ and Sr²⁺ differed, however. The lack of a concentrationgradient for Rb⁺ across the root was compatible with its transportmainly in the symplast, constituted by occasional files of intactcortical cells bridging the gas-filled spaces. The evident concentrationgradient for Sr²⁺ was accounted for by its movement to the endodermisby the apoplastic pathway constituted by the walls of intactand lysed cells. Key words: Aeration, aerenchyma, cations, electron probe X-ray microanalysis, Zea mays     

Measurements of mitochondrial K+ fluxes in whole rat hearts using 87Rb-NMR

The rubidium efflux from hypothermic rat hearts perfused by theLangendorff method at 20°C was studied. At thistemperature ⁸⁷Rb-NMR efflux experiments showed theexistence of two ⁸⁷Rb pools: cytoplasmic and mitochondrial.Rat heart mitochondria showed a very slow exchange of mitochondrialRb⁺ for cytoplasmic K⁺. After washout ofcytosolic Rb⁺, mitochondria kept a stable Rb⁺level for >30 min. Rb⁺ efflux from mitochondria wasstimulated with 0.1 mM 2,4-dinitrophenol (DNP), by sarcolemmalpermeabilization and concomitant cellular energy depletion by saponin(0.01 mg/ml for 4 min) in the presence of a perfusate mimickingintracellular conditions, or by ATP-sensitive K (K_ATP)channel openers. DNP, a mitochondrial uncoupler, caused the onset ofmitochondrial Rb⁺ exchange; however, the washout was notcomplete (80 vs. 56% in control). Energy deprivation by saponin, whichpermeabilizes the sarcolemma, resulted in a rapid and completeRb⁺ efflux. The mitochondrial Rb⁺ efflux rateconstant (k) decreased in the presence of glibenclamide, aK_ATP channel inhibitor (5 µM;k = 0.204 ± 0.065 min¹; n = 8),or in the presence of ATP plus phosphocreatine (1.0 and 5.0 mM,respectively; k = 0.134 ± 0.021 min¹;n = 4) in the saponin experiments (saponin only;k = 0.321 ± 0.079 min¹; n = 3),indicating the inhibition of mitochondrial K_ATP channels. Thus hypothermia in combination with ⁸⁷Rb-NMR allowed theprobing of the mitochondrial K⁺ pool in whole heartswithout mitochondrial isolation.

     

Shoot-Dependent Regulation of Sodium and Potassium Fluxes in Roots of Whole Barley Seedlings

Unidirectional fluxes and the cytoplasmic and vacuolar contentsof potassium and sodium in root cells of intact barley seedlings(Hordeum vulgare L., cv. Villa) were determined by use of compartmentalanalysis. In addition, the net vacuolar accumulation J_cv andthe xylem transport ø_cx of K⁺ and Na⁺ were measured.Both of these data were needed for the evaluation of the effluxdata. Fluxes and compartmental contents of K⁺ and Na⁺ were comparableto data obtained with excised roots. The effect of the shoot-to-rootratio—as varied by partial excision of the seedlings seminalroots—on the fluxes and contents was investigated. Highershoot-to-root ratios induced an increase in xylem transport,in plasmalemma influx, and also in the cytoplasmic content ofK⁺ and Na⁺. With potassium the plasmalemma efflux was almostunaltered while the tonoplast fluxes and vacuolar content weredecreased (in presence of Na⁺). With sodium, on the other hand,the plasmalemma efflux and the tonoplast fluxes were also increasedin the plants having one root and a high shoot-to-root ratio.These changes occurred even under conditions of low humidity,when transpiration was low and guttation occurred. The latterwas also increased at the high shoot-to-root ratio. The observedchanges could be due to a relieved feedback control of ion fluxesby the shoot and mediated in part by a relatively higher supplyof photosynthates in the plants having one root In addition,hormonal signals were suggested to participate. In particulara possibly decreased level of cytokinins in the plants havingonly one root could contribute to the signal. The observed changesappear to be responses of the plant to an alteration that canoccur under natural conditions when the root system is damaged.     

Inhibition of Ion Transport in Excised Barley Roots by Abscisic Acid; Relation to Water Permeability of the Roots

Previous papers have shown that abscisic acid can inhibit transportof ions across the root to the xylem vessels, resulting in reducedexudation from excised roots or inhibiting guttation from intactplants. However, it has not been established whether the inhibitionwas due to a reduction in salt transport (J_s) or in permeabilityof the roots to water (L_p). This paper investigates the effectof ABA on L_p and J_s separately. It is shown that L_p increasedin ABA and then fell, but was about the same as in control rootswhen transport was inhibited. The effect of ABA on exudationtherefore appeared to be mainly due to reduction in J_s. Inhibitionof J_s was also present in intact, transpiring plants and sowas not due to reduced water flow. The inhibition of ion releaseto the xylem affected Na⁺, Mg²⁺, Ca²⁺, and phosphate as wellas the major ion in the exudate, K⁺. It is concluded that ABAinhibits salt transport to the shoot by acting on ion transportinto the xylem, and not by reducing water flow coupled withsalt transport.     

Abnormal Stomatal Behaviour and Ion Imbalance in Capsicum scabrous diminutive

An attempt was made to explain the abnormal behaviour of stomatain Capsicum scabrous diminutive, a wilty pepper mutant. Stomatalmovement in the pepper plant was found to be associated withchanges in the ion content of the guard cells. These changeswere smaller in the mutant than in the normal plants. In addition,total ion content was higher in the mutant under both controland NaCl treatments. Na⁺ substituted K⁺ in its function in stomatalmovement under high salinity. This phenomenon was more pronouncedin the mutant plants. Analysis of whole root systems and leavesof plants grown on solutions of high NaCl or KCl concentrationconfirmed that the regulation of K⁺ and Na⁺ uptake mechanismswas not functioning properly in the mutant. Evidence was presentedthat the difference in K⁺ staining between mutant and normalepidermal cells is an artefact resulting from the differencein leaf anatomy.     

Potassium Transport in Non-Growing Corn Root Tissue as Affected by IAA and GA3

Experiments were done to determine if the spontaneous recoveryof non-growing segments of corn root (Zea mays L.) from excisioninjury is dependent on auxin. Washing the segments with 5 runindoleacetic acid (IAA) for 2 to 4 hours gave a small but significantincrease in K⁺ (⁸⁶Rb) influx, used here as a parameter reflectingrecovery of electrogenie H⁺-efflux pumping. This promotive effectwas obtained only after an hour of washing, and was sustainedby 100 nm gibberellic acid (GA₃). Any early responses to auxinwere obscured by an adverse reaction of the root cells to externalIAA which resulted in a transitory inhibition of H⁺ pumpingand K⁺ influx. Pretreatment of excised root tips with 10 µM IAA in thegrinding medium protected a plasmalemma-enriched fraction ofthe microsomes during isolation, giving increased uncoupler-sensitiveATPase activity. Non-growing root tissue thus shows three responses to auxin:an adverse reaction at the outer surface of the plasmalemmawhich blocks H⁺ pumping; a protective or restorative effecton the H⁺-ATPase; an increased capacity for K⁺ influx duringthe developmental phase of washing, which is augmented by thepresence of GA₃. (Received March 31, 1986; Accepted September 8, 1986)     

Permeation of Ca2+ and monovalent cations through an outwardly rectifying channel in maize root stelar cells

A depolarization-activated outwardly-rectifying channel (OR),most likely involved in the passive release of K⁺ from the rootsymplasm into the stelar apoplast (for subsequent transportto the shoot via the xylem vessels), has been characterizedin the plasma membrane of maize root stelar cells (Roberts andTester, 1995). In the present study, the selectivity of thischannel was further characterized using single channel current-voltagecurves generated using a voltage ramp protocol. This protocolpermitted the accurate and unambiguous measurement of the reversalpotentials of currents resulting from single channel openings.Using the voltage ramp protocol, it was shown that the OR allowsboth K⁺ efflux and Ca²⁺ influx at potentials positive of E_Kand negative of E_Ca. The OR had a P_Ca/P_K of 1.72–0.21decreasing as extracellular Ca²⁺ was increased. The permeabilityof the OR for monovalent cations other than K⁺ was also investigated.In biionic conditions, a relative permeability sequence of was determined (i.e. Eisenman sequenceIV). The physiological implications of the selectivity of theOR are discussed. Key words: Maize roots, K⁺ channel selectivity, Ca²⁺ permeation