Comparative Studies of Leaf Tissue and Isolated Mesophyll Protoplasts: II. ION RELATIONS

Freshly isolated tobacco mesophyll protoplasts had contentsof K^$, Cl^– and Na^$ slightly higher (on a cell basis) thanthe original leaf tissue, and had a high K^$/Na^$ ratio similarto that of the leaf tissue. Influxes of these ions into theprotoplasts were of similar magnitudes to the correspondingfluxes in leaf tissue when compared on the basis of the respectiveplasmalemma surface areas. In both systems the K^$ influx wasstrongly inhibited by CN^– and by DNP. These results suggestthat the ion relations of the freshly isolated mesophyll protoplastswere similar to those of the original leaf tissue and that isolatedmesophyll protoplasts should be a useful system for the studyof ion transport processes in leaf cells.     

Effects of Fusicoccin and Abscisic Acid on Sugar and lon Transport from Plant Glands

Fusicoccin (FC) inhibited net excretion of Cl^– by theglands of the pitchers of the carnivorous plant Nepenthes hookeriana;of Na⁺ and Cl^– by the salt glands of the halophytes Limoniumvulgare and L. pectinatum and of K⁺ in the nectar of Acer platanoidesflowers. It had no effect on K⁺ elimination with nectar of Impatienswalleriana (extrafloral nectaries) and Abutilon striatum. Abscisicacid (ABA) stimulated net excretion of K⁺ and Cl^– in Nepenthesand of Na⁺ and Cl^– in Limonium but had no effects on K⁺in nectar. Thus, FC and ABA had opposing effects on ion excretionby the salt eliminating glands of Limonium and Nepenthes. Bothcompounds, however, had similar effects on sugar secretion ofnectary glands which was either inhibited or unaffected by FCand ABA. It is suggested that the effects of FC and ABA on ion excretionby gland cells could be reconciled with literature showing FC-stimulationand possible ABA-inhibition of proton pumps at the plasmalemmaof plant cells. Nepenthes hookeriana, Limonium vulgare, Limonium pectinatum, Acer platanoides, salt-glands, nectaries, excretion, fusicoccin, abscisic acid, proton pump     

The intercellular distribution of vacuolar solutes in the epidermis and mesophyll of barley leaves changes in response to NaCl

Concentrations of inorganic and organic solutes were measuredin sap extracted from individual mesophyll and epidermal cellsof the third leaf of barley. During the development of the thirdleaf plants were grown in various salt solutions (NaCl; 2, 50,100, and 150 mM, KCI; 100 mM or KNO₃; 100 mM). Leaves were analysed2–4 d after full expansion. Cell-sap was extracted usinga modified pressure probe and analysed for its osmolality, concentrationsof P, Na⁺ K⁺ Ca²⁺, and Cl^– and, in some cases, of nitrate,hexoses and total amino acids. Salt treatment caused differentialchanges in the concentrations of solutes in mesophyll and epidermalcells, but did not affect the basic pattern of solute compartmentationbetween these tissues. Calcium was found at osmotically significantconcentrations only in the epidermis, whereas P and organicsolutes were almost exclusively found in the mesophyll. Chlorideand Na⁺ accumulated preferentially in the epidermis, althoughmesophyll concentrations also increased considerably. At 150mM external NaCl, mesophyll cells contained 302 mM Na and 167mM Cl^–, compared to 29 mM Na⁺ and 16 mM Cl^– in thecontrol. Mesophyll Cl^– levels were even higher in the100 mM KCl treatment (216 mM) where mesophyll and epidermalK⁺ accumulated to 424 and 491 mM, respectively. These huge increasesin mesophyll Na⁺ Cl^– and K⁺ were not associated with abreakdown in leaf performance since net rates of photosynthesisdecreased only by less than 20%. Under control (2 mM NaCl) conditions,solutes followed patterned gradients between the various epidermalcell types. The extent of these gradients changed with leafage. During 50 mM NaCl treatment, gradients in Cl^–, nitrateand malate concentrations progressively disappeared, with malateconcentrations approaching zero. Potassium and Na⁺ exhibitedaltered distribution profiles, whereas Ca²⁺ distribution wasunaffected. NaCl-dependent increases in osmolalities differedbetween cells. Exposure of plants to 150 mM NaCl caused qualitativelysimilar changes in both epidermal solute and osmolality profiles,although absolute values differed from those at 50 mM NaCl.In particular, epidermal Cl^– and Na⁺ increased to about500 mM and K⁺ disappeared (<<5 mM) from the vacuole ofcertain epidermal cell types completely. Key words: Barley leaf epidermis, mesophyll, salt stress, single-cell analysis, vacuolar solutes     

Measurement of the Apoplastic Activity of K+ and Cl- in the Leaf Epidermis of Commelina communis in Relation to Stomatal Activity

Bowling, D. J. F. 1987. Measurement of the apoplastic activityof K⁺ and Cl^– in the leaf epidermis of Commelina communisin relation to stomatal activity.–J. exp. Bot. 38: 1351–1355. Ionic activity of K⁺ and Cl^– in the apoplast of the lowerepidermis of the leaf of Commelina communis was measured usingion selective micro-electrodes. Large gradients across the stomatalcomplex were observed which were related to stomatal aperture.On stomatal closure the activity of K⁺ and Cl^– in theapoplast of the guard cell rose from 3·0 mol m^–3to 100 mol m^–3 and 33 mol m^–3 respectively. It wasconcluded that the apoplast is an important pathway for iontransport between the cells. Key words: Stomata, ionic activity, leaves, apoplast     

Inhibition of Sugar and Salt Elimination by Glands with the Amino Acid Analog p-Fluorophenylalanine

The amino acid analog p-fluorophenylalanine (FPA) inhibitedsugar and K⁺ secretion by nectary glands. FPA specifically reducedthe net excretion of Na⁺ and Cl^– by the salt glands ofthe halophyte Limonium vulgare and ³⁶Cl^–excretion by theglands of the pitcher walls of the carnivorous plant Nepenthes.Net uptake and net accumulation of Na⁺ and Cl^– by Limoniumleaf tissue and ³⁶Cl^– accumulation in Nepenthes pitchertissue were much less inhibited than excretion. The resultsare discussed in relation to literature reporting similar specificeffects of FPA on transport of ions from the symplast of barleyroots into the dead xylem elements. Limonium vulgare, Nepenthes hookeriana, salt-glands, excretion, p-fluorophenylalanine     

Some Observations on the Effects of Salinity on Ion Distributions and Cell Ultrastructure in Wheat Leaf Mesophyll Cells

The distribution of ions within wheat leaf mesophyll cells fromplants grown in the presence of 100 mol m^–3 NaCl werestudied by X-ray micro-analysis, and the volume fractions occupiedby the analysed compartments were determined stereologically.Generally, the cells did not accumulate high ion concentrationsin their vacuoles, and the much higher Na⁺ and Cl^– concentrationsin the cell walls would produce an imbalance of cell water relations.The Na⁺, K⁺ and Cl^– concentrations in the cytoplasm wereoften too high to be compatible with biochemical activity. Inresponse to saline growth conditions, the volume fractions occupiedby the cytoplasm and chloroplasts increased. Correspondence to: Institut für Botanik, Fachbereich Biologie,Technische Hochschule Darmstadt, 3–5 Schnittspahnstr.F.R.G. Key words: Salinity sensitivity, X-ray micro-analysis, stereology     

Ionic Currents across the Plasmalemma of Chara inflata Cells: I. OSMOTIC EFFECTS OF SORBITOL ON K+, CI- AND LEAK CURRENTS

This paper describes experiments designed to investigate theeffects of increases in external osmotic pressure on the electrophysiologicalbehaviour of the plasmalemma in cells of the brackish-wateralga, Chara inflata. The electrical conductance of the plasmalemmaof these cells of, due to the diffusion of ions, consists mainlyof K⁺, Cl^– and leak components. The addition of sorbitolat concentrations in the range 40–280 mol m^–3 tothe external solution bathing the cells, progressively and reversiblydepolarized the cell membrane and increased the total membraneconductance, chiefly due to increases in each of the separateionic conductances. At concentrations higher than about 280mol m^–3 when the cells became plasmolysed, the effectsof sorbitol on the electrical properties of the cell ceasedto be fully reversible. When the membrane electrical potentialdifference is stepped in a negative direction with a voltage-clamp,the resulting inward current has voltage-dependent componentsconsisting of an inactivating K⁺ current, an activating Cl^–current and a constant leak current. The addition of sorbitolto the external solution modified these currents by increasingtheir magnitude, by increasing the half-time of the inactivationof the K⁺ current, and by decreasing the half-time of activationof the Cl^– current. Key words: Chara inflata, osmotic effects, K⁺ and Cl^– currents     

Diurnal K+ and Anion Transport in Phaseolus Pulvinus

Diurnal movement of Phaseolus leaf is caused by deformationof the laminar pulvinus located at the joint of the leaf bladeand the petiole. The plants were cultured in solutions withvarious ion compositions, and changes of K⁺, Na⁺, Ca²⁺, Mg²⁺,Cl^–, NO₃– and P₁ concentrations both in the upperand lower parts of the laminar pulvinus were measured. Culturein 10 mM KCl solution caused an increase in K⁺ and Cl^–concentrations both in the upper and lower parts without anysignificant change in the concentration of NO₃^–; culturein 10 mM KNO₃ solution caused an increase in K⁺ and NO₃^–concentration without any significant change in the concentrationof Cl^–; and culture in 10 mM KH₂PO₄ solution caused anincrease in K⁺ and P₁ concentrations without any significantchange in the concentrations of NO₃- and Cl^–. K⁺ moved from the upper to lower parts or from the lower toupper parts diurnally in all plants cultured in any solutionmentioned above. The main inorganic anion that accompanied thisK⁺ movement was Cl^– in KCl solution, and NO₃^– inKNO₃ solution. When the seedlings were cultured in distilledwater or in KH₂PO₄ solution, neither Cl^– NO₃^– norP₁ accompanied this K⁺ movement. In these cases, mainly H⁺ and/ororganic anions are supposed to move in exchange for and/or incombination with K⁺ movement. (Received November 8, 1982; Accepted June 13, 1983)     

Ion Effluxes during Excitation of Characeae

Ion effluxes during excitation of Chara and Nitellopsis measuredby conductometry method were compared with results obtainedby two other analytical methods, Ag-AgCl method for Cl^–and ion chromatography method for K⁺. In both species, the averageefflux measured by the conductometry method was very close tothose of K⁺ and Cl^–. (Received May 12, 1986; Accepted June 25, 1986)     

Cl- secretory effects of EBIO in the rabbit conjunctival epithelium

Experiments were conducted to determine whether the Cl^– secretagogue, 1-ethyl-2-benzimidazolinone (EBIO), stimulates Cl^– transport in the rabbit conjunctival epithelium. For this study, epithelia were isolated in an Ussing-type chamber under short-circuit conditions. The effects of EBIO on the short-circuit current (I_sc) and transepithelial resistance (R_t) were measured under physiological conditions, as well as in experiments with altered electrolyte concentrations. Addition of 0.5 mM EBIO to the apical bath stimulated the control I_sc by 64% and reduced R_t by 21% (P < 0.05; paired data). Under Cl^–-free conditions, I_sc stimulation using EBIO was markedly attenuated. In the presence of an apical-to-basolateral K⁺ gradient and permeabilization of the apical membrane, the majority of the I_sc reflected the transcellular movement of K⁺ via basolateral K⁺ channels. Under these conditions, EBIO in combination with A23187 elicited nearly instantaneous 60–90% increases in I_sc that were sensitive to the calmodulin antagonist calmidazolium and the K⁺ channel blocker tetraethyl ammonium. In the presence of an apical-to-basolateral Cl^– gradient and nystatin permeabilization of the basolateral aspect, EBIO increased the Cl^–-dependent I_sc, an effect prevented by the channel blocker glibenclamide (0.3 mM). The latter compound also was used to determine the proportion of EBIO-evoked unidirectional ³⁶Cl^– fluxes in the presence of the Cl^– gradient that traversed the epithelium transcellularly. Overall, EBIO activated apical Cl^– channels and basolateral K⁺ channels (presumably those that are Ca²⁺ dependent), thereby suggesting that this compound, or related derivatives, may be suitable as topical agents to stimulate fluid transport across the tissue in individuals with lacrimal gland deficiencies. Ussing chamber; short-circuit current; electrolyte transport; chloride secretagogue; potassium conductance; 1-ethyl-2-benzimidazolinone; 1,10-phenanthroline     

Biophysical and Biochemical Regulation of Cytoplasmic pH in Chara corallina during Acid Loads

The contribution of membrane transport to regulation of cytoplasmicpH in Chara corallina has been measured during proton-loadingby uptake of butyric acid. In the short-term (i.e. up to 20min) uptake of butyric acid is not affected by removal of externalK⁺, Na⁺ or Cl^– but over longer periods uptake is decreased(by 20–50% in different experiments) in the absence ofexternal Na⁺ or, sometimes, K⁺. Influxes of both Na⁺ and K⁺increase temporarily after addition of butyrate, Na⁺ immediatelyand K⁺ after a lag. Effects on Cl^– influx are small butCl^– efflux increases enormously after a short lag. Anapproximate comparison of internal butyrate with changes inthe concentration of K⁺, Na⁺, and Cl^– suggests that initially(i.e. for a few min) cytoplasmic pH is determined by bufferingand possibly by some decarboxylation of organic acids (biochemicalpH regulation), and that biophysical pH regulation involvingefflux of H⁺ balanced by influxes of K⁺, Na⁺ and especiallyefflux of Cl^– progressively becomes dominant. When butyric acid is washed out of the cells, cytoplasmic pHis restored completely or partially (depending on the butyrateconcentration used) and this is independent of the presenceor absence of external Cl^–. Where Cl^– is present,its influx is relatively small. It is suggested that cytoplasmicpH is then controlled biochemically, involving the synthesisof an (unidentified) organic acid and the accumulation of acidicanions in place of butyurate lost from the cell. During thesecond application of butyrate, net Cl^– efflux is small:it is suggested that control of cytoplasmic pH then involvesdecarboxylation of the organic acid anions. The questions of the source of Cl^– lost from the cell(cytoplasm or vacuole) and of possible cytoplasmic swellingassociated with the accumulation of butyrate are discussed. Key words: Chara corallina, butyric acid, cytoplasmic pH, membrane transport     

The Ionic Relations of Acetabularia mediterranea

The concentrations of K⁺, Na⁺, and Cl^– in the cytoplasmand the vacuole of Acetabularia mediterranea have been measured,as have the vacuolar concentrations of SO₄^–– andoxalate. The electrical potential difference between externalsolution, and vacuole and cytoplasm has been measured. The resultsindicate that Cl^– and SO₄^–– are probably transportedactively into the cell, and that active transport of Na⁺ isoutwards. The results for K⁺ are equivocal. The fluxes of K⁺,Na⁺, Cl^–, and S0₄^–– into the cell and theeffluxes of Na⁺ and Cl^– have been determined. The Cl^–fluxes are extremely large. In all cases the plasmalemma isthe rate-limiting membrane for ion movement. A technique isdescribed for the preparation of large, completely viable cellfragments containing only cytoplasm, with no vacuole.     

Ionic Activity Gradients across the Surface Membrane of Cytoplasmic Droplets Prepared from Chara australis

The membrane potential and the ionic activity gradients of K⁺and Cl^– across the surface membrane of cytoplasmic dropletsprepared from Chara australis internodal cells, were measuredin high and low ionic strength bathing solutions using liquidion exchange microelectrodes selective for K⁺ and Cl^–.Our results indicate that K⁺ is close to electrochemical equilibriumwhereas Cl^– is not. ¹ Present address: ICI Japan, Palace Hotel Annex Building, Marunouchi,Tokyo, Japan.     

Protective role of extracellular chloride in fatigue of isolated mammalian skeletal muscle

A possible role of extracellular Cl^– concentration ([Cl^–]_o) in fatigue was investigated in isolated skeletal muscles of the mouse. When [Cl^–]_o was lowered from 128 to 10 mM, peak tetanic force was unchanged, fade was exacerbated (wire stimulation electrodes), and a hump appeared during tetanic relaxation in both nonfatigued slow-twitch soleus and fast-twitch extensor digitorum longus (EDL) muscles. Low [Cl^–]_o increased the rate of fatigue 1) with prolonged, continuous tetanic stimulation in soleus, 2) with repeated intermittent tetanic stimulation in soleus or EDL, and 3) to a greater extent with repeated tetanic stimulation when wire stimulation electrodes were used rather than plate stimulation electrodes in soleus. In nonfatigued soleus muscles, application of 9 mM K⁺ with low [Cl^–]_o caused more rapid and greater tetanic force depression, along with greater depolarization, than was evident at normal [Cl^–]_o. These effects of raised [K⁺]_o and low [Cl^–]_o were synergistic. From these data, we suggest that normal [Cl^–]_o provides protection against fatigue involving high-intensity contractions in both fast- and slow-twitch mammalian muscle. This phenomenon possibly involves attenuation of the depolarization caused by stimulation- or exercise-induced run-down of the transsarcolemmal K⁺ gradient. potassium; skeletal muscle contraction; membrane potential; myotonia     

Ion Fluxes Across the Transfusion Zone of Secreting Limonium Salt Glands Determined from Secretion Rates, Transfusion Zone Areas and Plasmodesmatal Frequencies

Faraday, C. D., Quinton, P. M. and Thomson, W. W. 1986. Ionfluxes across the transfusion zone of secreting Limonium saltglands determined from secretion rates, transfusion zone areasand plasmodesmatal frequencies.—J. exp. Bot. 37: 482–494. The epidermal salt-secreting glands of Limonium (Plumbaginaceae)are enclosed in a cuticular envelope. Ions and metabolites enterthe glands from the mesophyll through gaps in the cuticularenvelope, the transfusion zones. Net influxes of ions acrossthe transfusion zone were calculated from measurements of secretionrates and transfusion zone areas. When leaves of L. pereziiF. T. Hubb. were treated with 300 mol m^–3 NaCl, transfusionzone influxes of Na⁺ K⁺, Ca⁺⁺ and Cl^– as high as 7?0?10^–5,1.7?10^–5, 5?8?10^–7 and 8.5?10^–5 mol m^–2s^–1 respectively, were calculated. Assuming a transmembranepathway, these fluxes would be some of the highest reportedfor ions in plant cells. Key words: Salt glands, ion fluxes, ultrastructure     

Separation of K+-and Cl --selective ion channels from rye roots on a continuous sucrose density gradient

Microsomal membranes from rye (Secale cereale L.) roots wereseparated by isopycnic sucrose density gradient centrifugation.The ion channels present in gradient fractions were assayedby reconstitution into planar 1-palmitoyl-2-oleoyl phosphatidylethanolaminebilayers (PLB) and the distributions of ion channel activitieswere compared with membrane markerenzyme activities. A numberof ion channel activities were observed and could be distinguishedon the combined bases of their conductance, selectivity, kineticsand pharmacology. A voltage-dependent maxi (498 pS) cation-channel,a voltage-dependent 199-pS cationchannel, 48-pS and 18-pS K⁺channels, and a 148-pS Cl^– channel (all unitary conductancesdetermined in asymmetrical cis trans 325:100mM KCl) colocalizedwith the plasma membrane marker-enzyme, vanadatesensitive ATPase.A weakly K ⁺-selective (108 pS) channel, a 1249-pS cation-channeland a 98-pS K ⁺ channel colocalized with the tonoplast markerenzyme,nitrate-sensitive ATPase. A 706-pS K⁺ channel colocalized withthe expected distribution of intact plastids and a 38-pS Cl^–channel colocalized with either plastid or ER membranes. Themembrane location of several other channels including a hypervoltage-sensitivemaxi (497 pS) cation-channel, a 270-pS K⁺ channel, an 8-pS K⁺channel and a 4-pS K⁺ channel was equivocal, but they were tentativelyassigned to the Golgi. Thus, the plasma membrane and tonoplastorigin of ion channels previously characterized following theincorporation of plasma membrane prepared by aqueous-polymertwo-phase partitioning or tonoplast derived from isolated vacuolesinto PLB was confirmed and the ion channel complement of previouslyunassayed membranes was defined. This demonstrates the usefulnessof PLB in identifying and characterizing ion channels from plantcell membranes, in particular, those of membranes which areinaccessible to patch-clamp electrodes. Key words: Chloride (Cl^–) channel, potassium (K⁺) channel, planar lipid bilayer, root, rye, Secale cerealeL.     

The Role of Transmembrane Electrical Potential in Determining the Absorption Isotherm for Chloride in Potato

Measurements have been made of the electrical potential differencebetween the vacuoles of single potato tuber cells and externalCl^- solutions over the range 1–40 mM. With K⁺ as the counter-ion,the relationship between this transmembrane electrical potentialand external Cl^- concentration, for fresh cells at 20° C,was found to be one of decreasing negative polarity with increasingCl^- concentration (E at 1 mM Cl^- external = – 81 m V;change in E for a 10-fold change in external concentration,E₁₀ = 46 m V). The linearity of this relationship, apparenton a semi-logarithmic plot, was virtually unaltered by low temperature(0.5–2.5° C) or by previous ageing of the cells forperiods up to four days (indicating that metabolic ion absorptionis not an electrogenic process). When the counter-ion was maintainedat a constant high concentration (40 mM K⁺), the change in potentialover the Cl^- concentration range was only 4 m V, polarity becomingmore negative with increasing Cl^- concentration. With Ca⁺⁺ asthe counter-ion, the potential to external Cl^- concentrationrelationship was similar to that found in KCl solutions, exceptthat E₁₀ was only about 20 m V. Curves for the influx of C1^- to be expected on the basis ofthese electrochemical data alone have been shown to run closelyparallel to Cl^– absorption isotherms previously determinedexperimentally. This confirms the opinion, already formed onthe basis of theoretically derived values for passive Cl^- influx,that Cl^- uptake by both fresh and one-day-aged potato tissue,from KCl solutions and Cl^- solutions with a fixed high K⁺ concentration,is rate-determined at o° C by passive movement across theplasmalemma. Uptake of Cl^- by fresh tissue at 20° C appearsto be similarly regulated. No such parallelism was found between observed and expectedpatterns of Cl^- uptake from CaCl₂ solutions, or from KCl bytwo-day-aged tissue, and here factors in addition to the electrochemicalones must determine low temperature Cl^-uptake.     

Involvement of anion channel(s) in the modulation of the transient outward K(+) channel in rat ventricular myocytes

The cardiac Ca²⁺-independent transient outward K⁺ current (I_to), a major repolarizing ionic current, is markedly affected by Cl^– substitution and anion channel blockers. We reexplored the mechanism of the action of anions on I_to by using whole cell patch-clamp in single isolated rat cardiac ventricular myocytes. The transient outward current was sensitive to blockade by 4-aminopyridine (4-AP) and was abolished by Cs⁺ substitution for intracellular K⁺. Replacement of most of the extracellular Cl^– with less permeant anions, aspartate (Asp^–) and glutamate (Glu^–), markedly suppressed the current. Removal of external Na⁺ or stabilization of F-actin with phalloidin did not significantly affect the inhibitory action of less permeant anions on I_to. In contrast, the permeant Cl^– substitute Br^– did not markedly affect the current, whereas F^– substitution for Cl^– induced a slight inhibition. The I_to elicited during Br^– substitution for Cl^– was also sensitive to blockade by 4-AP. The ability of Cl^– substitutes to induce rightward shifts of the steady-state inactivation curve of I_to was in the following sequence: NO₃^– > Cl^– Br^– > gluconate^– > Glu^– > Asp^–. Depolymerization of actin filaments with cytochalasin D (CytD) induced an effect on the steady-state inactivation of I_to similar to that of less permeant anions. Fluorescent phalloidin staining experiments revealed that CytD-pretreatment significantly decreased the intensity of FITC-phalloidin staining of F-actin, whereas Asp^– substitution for Cl^– was without significant effect on the intensity. These results suggest that the I_to channel is modulated by anion channel(s), in which the actin cytoskeleton may be implicated. transient outward potassium current; anion channel; actin cytoskeleton; myocyte; potassium ion     

Chloride Transport in Chara: I. KINETICS AND CURRENT-VOLTAGE CURVES FOR A PROBABLE PROTON SYMPORT

Chara cells show an inward positive electric current acrossthe plasmalemma when exposed to Cl^– under voltage-clampconditions. The rapid rise of this current suggests that itis directly associated with the inward transport of Cl^–.The dependence of the current on Cl^– concentration showssaturation, the data fitting the Michaelis-Menten equation withV_m up to 100 nmol m^–2 s^–1 (for Cl^–starvedcells) with K_M 10–20 µM, and with some allowancefor an unstirred layer of water adjacent to the membrane. Theeffects on the current of clamp potential, illumination, withdrawalof alkali metal cations, and addition of amine were also investigated.These results suggest that the mechanism is the symport of 2H⁺ with each Cl^–, and that the actions of light, externalK⁺, and amine in stimulating Cl^–, influx are indirect.     

Electrophysiological Measurements on the Root of Atriplex hastata

The ion contents and membrane potentials of the cells of young,hydroponically cultured seedlings of Atriplex hastata L. var.salina, Wallr. have been measured at several different NaClconcentrations. The total tissue concentrations of Na⁺ and Cl^–increase as external NaCl increases, but there is always a markedexcess of internal Na⁺ over Cl^–; this is balanced by endogenousorganic anion formation with a concomitant extrusion of H⁺ tothe bathing solution. Membrane potentials of the root cells remain essentially invariantwith changes in external NaCl at approx. –130 mV; thereis no evidence of a radial gradient of potential across theroot. The potential seems to contain a cyanide-sensitive electrogeniccomponent, also invariant with NaCl concentration, of about–70 mV, and a diffusion component. The electrogenic componentseems likely to be a H⁺ efflux, probably through a H⁺ uniportATPase.