Interaction of Salinity and Anaerobiosis in Barley and Rice

Barley and rice at the early tillering stage were exposed simultaneouslyto anaerobiosis and high [NaCl]. Barley was grown at 0.5, 70,and 125 mol m^–3 NaCl, and rice at 2, 20, 40, and 80 molm^–3 NaCl. Surprisingly, anaerobiosis only slightly aggravatedthe adverse effects of high [NaCl] on root and shoot growthof both species. For rice and barley grown under aerobic conditions, high [NaCl]increased [Na⁺] and [Cl^–] and decreased [K+] in both rootsand shoots. However, the changes in ion concentrations in theshoots were smaller for rice than for barley. For roots of barley, anaerobiosis decreased [Na⁺], [Cl^–],and [K⁺] at both low and high [NaCl], possibly as a result ofinhibition of active ion accumulation. For barley shoots, anaerobiosisincreased [Na⁺] and [Cl^–], but only at high salinity;in contrast, [K⁺] was reduced by anaerobiosis at both low andhigh [NaCl]. These results indicate that anaerobiosis slightlyincreased the permeability of the barley root system to Na+and Cl^–. For rice, the most important interaction between salinity andanaerobiosis occurred in the shoots, where anaerobiosis increased[Na⁺] and decreased [K⁺], particularly at 40 and 80 mol m^–3NaCl, while there was no interaction between anaerobiosis andsalinity for Cl^– uptake. It is therefore suggested thatanaerobic treatment of rice decreased the selectivity for K⁺over Na⁺ of cation transport to the shoots, at least for plantsgrown at high salinities.     

Na+, K+ and Cl- in Xylem Sap Flowing to Shoots of NaCl-Treated Barley

Munns, R. 1985. Na⁺, K⁺ and Cl^– in xylem sap flowing toshoots of NaCl-treated barley.—J. exp. Bot. 36: 1032–1042. Na⁺, Cl^– and K⁺ concentrations were measured in xylemsap obtained by applying pressure to the roots of decapitatedbarley plants grown at external [NaCl] of 0, 25, 50, 100, 150and 200 mol m^–3. For any given NaCl treatment, ion concentrationsin the xylem sap were hyperbolically related to the flux ofwater. Ion concentrations in sap collected at very low volumefluxes (without applied pressure) were 5–10 times higherthan in sap collected at moderate fluxes (under pressure). Fora given moderate volume flux, Na+ concentration in the xylemsap, [Na⁺]_x, was only 4.0 mol m^–3 at external [NaCl] of25–150 mol m^–3, and increased to 7.0 mol m^–3at 200 mol m^–3. [Cl-]x showed a similar pattern. Thisshows there would be little difference in the rate of uptaketo the shoot of plants at 25–150 mol m^–3 externalNaCl and indicates little change even at 200 mol m^-3 NaCl becausetranspiration rates would be much lower. Thus the reduced growthof the shoot of plants at high NaCl concentrations is not dueto higher uptake rates of Na⁺ or Cl^–. The fluxes of Na⁺, Cl^– and K^– increased non-linearlywith increasing volume flux indicating little movement of saltin the apoplast. The flux of K⁺ increased even when [K⁺]_x wasgreater than external [K⁺], indicating that membrane transportprocesses modify the K⁺ concentration in the transpiration streamas it flows through the root system. Key words: -Xylem sap, Na⁺, K⁺, Cl^– fluxes, salinity, barley     

Effects of Sodium Chloride Stress on Callus Cultures of Cicer arietinum L. cv. BG-203: Growth and Ion Accumulation

Growth and ion accumulation were measured in callus culturesof Cicer arietinum L. cv. BG-203, grown on media supplementedwith 0–200 mol m^–3 NaCl. Fresh and dry weights decreasedat concentrations ranging from 100–200 mol m^–3,the reduction being greater during the third and fourth weeksof culture. Slight stimulation of growth was observed at 25and 50 mol m^–3 NaCl. There was also a decrease in tissuewater content (fresh weight: dry weight) at 100–200 molm^–3 NaCl. The concentration of Na⁺ and Cl^– in thetissue increased with increasing salinity of the medium. Mostof the accumulation of these ions occurred by the first weekwhile significant growth inhibition became apparent by onlythe third week of culture. Tissue K⁺ and Mg²⁺ decreased withincreasing salinization, the decrease being greater in K⁺ levels.Levels of Ca²⁺, however, were maintained throughout the experimentalrange. Key words: Cicer arietinum, NaCl stress, Callus cultures, Ion accumulation     

Effect of Salt Stress on Viability, Germination and Endogenous Levels of Some Metabolites and Ions in Maize (Zea mays L.) Pollen

Maize plants, subjected to 0, 80, 120 and 160 meq l^–1salinity using NaCl, showed adverse effects on viability, germinationand tube growth of pollen, besides enhancing the bursting ofpollen. The endogenous levels of various metabolites in pollenwere also affected. Pollen grains from salinized plants hadmore soluble carbohydrates, free amino acids, especially proline,phenols and DNA and less starch, protein and RNA compared tothe non-saline controls. Salinity also resulted in the accumulationof ions such as Na⁺, K⁺ and Cl^– while it caused a reductionin the boron content of pollen. These metabolic disturbancespossibly lead to decreased viability, germination and tube growthof pollen thereby resulting into a reduction in reproductivecapacity of the plants under salt stress. Zea mays L., maize, pollen, viability, germination, salt stress     

Effect of NaCl Salinity on Somatic Embryo Development in Sapindus trifoliatus L.

The effect of NaCl salinity on growth and development of somaticembryos of Sapindus trifoliatus L. was examined. Incorporationof 25 and 50 mol m^–3 NaCl into the medium greatly increasedthe growth and development of somatic embryos and both theseconcentrations favoured the production of secondary embryoids.However, supplementation of 100 mol m^–3 NaCl to the mediumdid not have any significant effect on the growth and developmentof somatic embryos. On the other hand, the culturing of proembryostructures in medium containing 200 mol m^–3 NaCl resultedin complete death within 7 d of salt exposure. Analysis of somatic embryos revealed that, upon salinization,they accumulated Na⁺ and Cl^– in significant amounts butthe content of Na⁺ was much less compared to that of Cl^–.Addition of NaCl (up to 50 mol m^–3) in the medium resultedin a considerable increase in the K⁺ content of somatic embryos.The content of proline in somatic embryos, however, increasedsubstantially in response to salinization. The amount of freesterols, steryl glycosides, steryl esters, and phospholipidsalso rose to higher values in salt-affected somatic embryos.The results suggest that somatic embryos of S. trifoliatus cantolerate concentrations of NaCl up to 100 mol m^–3 withoutaffecting growth and that they have sufficient cellular mechanismsto tolerate salinity at relatively high levels. Key words: Salinity, somatic embryo, sterols, phospholipids     

Salt Tolerance in the Succulent, Coastal Halophyte, Sarcocornia natalensis

The effects of 0, 50, 100, 200, 300, 400 and 500 mol m^–3NaCl on growth and ion accumulation in the succulent, coastalhalophyte Sarcocornia natalensis (Bunge ex Ung.-Sternb.) A.J. Scott were investigated. Increase in salinity from 0 to 300 mol m^–3 NaCl stimulatedproduction of fresh, dry, and organic dry mass, increased succulenceand shifted resource allocation from roots to shoots. Growthwas optimal at 300 mol m^–3 and decreased with furtherincrease in salinity. Water contributed to a large proportion of the increase in freshmass. Inorganic ions, especially Na⁺ and Cl– contributedsubstantially to the dry mass. At 300 mol m^–3 NaCl inorganicions contributed to 37% of total dry mass and NaCl concentrationin the shoots was 482 mol m^–3. Expressed sap osmotic potentialsdecreased from –2.10 to –3.95 MPa as salinity increasedfrom 0 to 300 mol m^–3 NaCl. Massive accumulation of inorganicions, especially Na⁺ and Cl^–, accounted for 86% of theosmotic adjustment at 300 mol m^–3 NaCl. Salinity treatments decreased the concentrations of K+ in shoots.Plant Na+ :K+ ratios increased steadily with salinity and reacheda maximum of 16.6 at 400 mol m3 NaCl. It is suggested that the exceptional salt tolerance of S. natalensisis achieved by massive inorganic ion accumulation which providessufficient solutes for osmoregulation, increased water fluxand turgor-induced growth. Key words: Sarcocornia natalensis, salt tolerance, halophyte     

The Effects of Salinity upon Ion Content and Ion Transport of the Marine RedAlga Porphyra purpurea(Roth) C.Ag.

Ion contents and concentrations (K⁺, Na⁺, Cl^–, Ca²⁺, Mg²⁺,SO^2–₄, NO^–₃, HPOJ^2–₄, amino and organic acids)of P. purpurea have been studied in relation to salinity variation.Cells were shown to accumulate large amounts of K⁺ and Cl^–against their respective gradients of electrochemical potentialin all dilute and concentrated seawater media. Active influxof SOJ^2–₄, NO^–₃, and HPOJ^2–₄ is also suggested,while Na⁺ is actively excluded from cells under hyposaline andhypersaline conditions. The relative proportions of individualcomponents of the internal osmotic potential were found to changeaccording to the external salt concentration. KCL forms themajor fraction of _j} in concentrated seawater media while K⁺-aminoacids form the major fraction in dilute seawaters. Other intracellularsolutes comprise less than 15% oft_j, in all media. Unidirectional fluxes of K⁺ and Cl^– were studied by radioisotopicmeans. Fluxes of K⁺ and Cl^– are reduced in hyposalinemedia, as is absolute KCL content per cell. Intracelhilar KCLcontent was also found to be markedly dependent upon externalK⁺ concentration, rather than water potential. Changes in KC1levels induced by salinity variation occur over a 6 h period.     

Measurements of Ion Concentrations inDunaliella parvaSubjected to Hypertonic Shock

Cells of Dunaliella parva, a green halotolerant alga, were equilibratedin 0.5 M NaCl (the same concentration as in the growth medium).The time – course of changes in K⁺, Na⁺, Cl^–, andpellet volume were followed after a hypertonic shock causedby increasing the outside NaCl concentration to 1 M. The cellsresponded initially by shrinking; they returned to their originalvolume in the course of the next 2–3 h. At the time ofthe hypertonic shock there were rapid influxes of Na⁺ and Cl^–which took about 30 min for completion. Thereafter, there wereprogressive losses of Na⁺ and Cl^–; these losses were temperature-dependentand presumably activated by metabolic energy. Cell K⁺ remainedconstant throughout.     

Cation and Chloride Partitioning through Xylem and Phloem within the Whole Plant of Ricinus communis L. under Conditions of Salt Stress

Uptake and partitioning through the xylem and phloem of K⁺,Na⁺, Mg²⁺ , Ca²⁺ and Cl^– were studied over a 9 d intervalduring late vegetative growth of castor bean (Ricinus communisL.) plants exposed to a mean salinity stress of 128 mol m^–3NaCl. Empirically based models of flow and utilization of eachion within the whole plant were constructed using informationon ion increments of plant parts, molar ratios of ions to carbonin phloem sap sampled from petioles and stem internodes andpreviously derived information on carbon flow between plantsparts in xylem and phloem in identical plant material. Salientfeatures of the plant budget for K⁺ were prominent depositionin leaves, high mobility of K⁺ in phloem, high rates of cyclingthrough leaves and downward translocation of K⁺ providing theroot with a large excess of K⁺ . Corresponding data for Na⁺showed marked retention in the root, lateral uptake from xylemby hypocotyl, stem internodes and petioles leading to low intakeby young leaf laminae and substantial cycling from older leavesback to the root. The partitioning of the anionic componentof NaCl salinity, Cl^–, contrasted to that of Na⁺ in thatit was not substantially retained in the root, but depositedmore or less uniformly in stem, petiole and leaf lamina tissues.The flow pattern for Mg²⁺ showed relatively even depositionthrough the plant but some preferential uptake by young leaves,generally lesser export than import by leaf laminae, and a returnflow of Mg²⁺ from shoot to root considerably less than the recordedincrement of the root. Ca²⁺ partitioning contrasted with thatof the other ions in showing extremely poor phloem mobility,leading to progressive preferential accumulation in leaf laminaeand negligible cycling of the element through leaves or root.Features of the response of Ricinus to salinity shown in thepresent study were discussed with data from similar modellingstudies on white lupin (Lupinus albus L.) and barley (Hordeumvulgare L.) Key words: Ricinus communis L, potassium, sodium, chloride, calcium, magnesium, phloem, xylem, transport, partitioning, salinity     

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     

Potassium Transport in Enteromorpha intestinalis (L.) Link: MEASUREMENT OF INTRACELLULAR K+, EXCHANGE FLUXES AND THERMODYNAMIC ANALYSIS

Potassium transport has been studied in the marine euryhalinealga, Enteromorpha intestimlis cultured in seawater and in low-salinitymedium (Artificial Cape Banks Spring Water, ACBSW; 25·5mol m^–3 Cl^–, 20·4 mol m^–3 Na⁺, 0·5mol m^–3 K⁺). K⁺ fluxes were measured using ⁴²K⁺ and ⁸⁶Rb⁺although ⁸⁶Rb⁺ does not act as an efficient K⁺ analogue in thisplant. ⁴²K⁺ experiments on seawater plants typically exhibiteda single protoplasmic exchange phase whereas ⁸⁶Rb⁺ exhibitedtwo exchange phases. Compartmental analysis of ⁸⁶Rb⁺ effluxexperiments on seawater-grown Enteromorpha plants were usedto deduce the intracellular partition of K⁺ between the cytoplasm(279±38 mMolal) and vacuole (405±68 mMolal). Theplasmalemma K⁺ flux in plants in seawater was greater in thelight than in the dark (563±108 nmol m^–2 s^–1versus 389±66·7 nmol m^–2 s^–1). Inlow-salinity plants, separate cytoplasmic and vacuolar exchangephases were apparent. Analysis of ⁴²K⁺ efflux experiments onlow-salinity plants yielded a cytoplasmic K⁺ of 222±38mMolal and a vacuolar K⁺ of 82±11 mMolal. The plasmalemmaand tonoplast flux was 23±4·5 nmol m^–2 s^–1. The Nernst equation showed that, although K⁺ was close to electrochemicalequilibrium, active accumulation of K⁺ across the plasmalemmaoccurred in plants in seawater and ACBSW both in the light anddark. K⁺ was also actively transported inwards across the tonoplastin low-salinity plants. The electrochemical potential for K⁺across the plasmalemma ranged from 2·41±0·60kJ mol^–1 in plants grown in seawater in the light to 5·79±0·87kJ mol^–1 for plants in ACBSW in the light. Although K⁺is close to electrochemical equilibrium, the flux of K⁺ in plantsin both seawater and ACBSW media is high, hence the power consumptionof K⁺ transport is high. The permeability of K⁺ (P_K⁺) was significantlyhigher in the light than in the dark in plants in seawater (about7·0 versus 2·5 nm s^–1) but in plants inlow-salinity (ACBSW) medium the permeability was independentof light (about 12 nm s^–1). The energy requirements ofactive K⁺ transport by ATP-dependent pumps is discussed. Key words: Enteromorpha, Potassium transport, Ionic relations, Saltwater, Low salinity, Thermodynamics     

Exogenous nitric oxide improves seed germination in wheat against mitochondrial oxidative damage induced by high salinity

Effects of exogenous nitric oxide (NO) on starch degradation, oxidation in mitochondria and K⁺/Na⁺ accumulation during seed germination of wheat were investigated under a high salinity level. Seeds of winter wheat (Triticum aestivum L., cv. Huaimai 17) were pre-soaked with 0 mM or 0.1 mM of sodium nitroprusside (SNP, as nitric oxide donor) for 20 h just before germination under 300 mM NaCl. At 300 mM NaCl, exogenous NO increased germination rate and weights of coleoptile and radicle, but decreased seed weight. Exogenous NO also enhanced seed respiration rate and ATP synthesis. In addition, seed starch content decreased while soluble sugar content increased by exogenous NO pre-treatment, which was in accordance with the improved amylase activities in the germinating seeds. Exogenous NO increased the activities of superoxide dismutase (SOD, EC 1.15.1.1) and catalase (CAT, EC 1.11.1.6); whereas decreased the contents of malondialdehyde (MDA) and hydrogen peroxide (H₂O₂), and superoxide anions (O₂^??) release rate in the mitochondria. Exogenous NO also decreased Na⁺ concentration while increased K⁺ concentration in the seeds thereby maintained a balance between K⁺ and Na⁺ during germination under salt stress. It is concluded that exogenous NO treatment on wheat seeds may be a good option to improve seed germination and crop establishment under saline conditions.     

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     

Induction of Increased Salt Tolerance in Sorghum bicolor by NaCl Pretreatment

Following 20 d of exposure to 75 or 150 mol m^–3 NaCl Sorghumbicolor (L.) Moench plants become capable of growing in mediumcontaining 300 mol m^–3 NaCl. Control plants, which havenot been pretreated, or plants pretreated for less than 20 ddie within 2 weeks when exposed to 300 mol m^–3 NaCl. Weconsider this induction of a capacity to survive in and toleratea high NaCl concentration as an adaptation to salinity. We suggestthat adaptation to salinity is more than osmotic adjustmentand that it takes longer to develop than osmotic adjustment.Concomitantly with the appearance of the ability to grow inhigh salinity, adaptation also comprises the development ofa capacity to regulate internal Na⁺ and Cl^– concentrations,even when external salinity is high. Shoot mean relative growthrates are similar for both control plants and for adapted plantsgrowing in 300 mol m^–3 NaCl, although their shoot Na⁺and Cl^– concentrations are quite different. Based on thesedata, we propose that adaptation of Sorghum to high salinityresults from a modulation of genome expression occurring duringextended exposure to non-lethal NaCl concentrations. Key words: Sorghum bicolor (L.) Moench, NaCl, salt tolerance, adaptation to salinity     

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     

Salinity-induced Malate Accumulation in Chara

Ion absorption by Chara corallina from solutions containingpredominantly KC1 or RbCl at up to 100 mol m^–3 resultedin accumulation of salts and turgor regulation. Turgor regulationdid not occur in solutions containing Na⁺ or Li⁺salts. Duringion absorption from various salts of K⁺ and Rb⁺ vacuolar cationconcentration exceeded Cl^– concentration. This differencewas shown to be balanced by the synthesis and accumulation ofmalate. Vacuolar malate concentration reached 48 mol m₃^–,with accumulation occurring at rates of up to 0.45 mol m^–3h^–1. Malate accumulation was inhibited by low externalpH and was dependent upon external HCO₃^– concentration.The synthesis of malic acid and its subsequent dissociationimposed a severe acid load on the cell. Biophysical regulationof cellular pH was achieved by a H⁺efflux at a rate of about40 nmol m^–2 s^–1from the cell. The results presentedargue against cytoplasmic Cl^–, HCO₃^– or pH regulatingmalate accumulation in Chara and it is suggested that malatetransport across the tonoplast may regulate malate accumulation. Key words: Malate, Chara corallina, pH regulation, salinity     

Effects of Anoxia on Solute Loss from Beetroot Storage Tissue

Beetroot storage tissue that had been aged in an aerated solutionwas particularly suited for studies of solute losses duringanoxia;retention of betacyanin being a good indicator of tonoplastintegrity. During anoxia, loss of K⁺ was nearly always greater than thatof Na⁺ while Cl^– loss was intermediate. Supply of glucoseduringageing increased the tolerance of beetroot tissue to anoxia.In these tolerant tissues, there were three phases of soluteloss.During the first phase, losses of K⁺ and amino acids wererapid, presumably due to membrane depolarization from –156to –95 mV. In contrast, losses of Na⁺ and Cl^– wereslow. During the second phase, K⁺ loss had decreased to a lowrate, while losses of Na⁺ and Cl⁺ remained slow. Furthermore,the membrane potential remained at –95 to –90mV,which was consistent with the diffusion potential estimatedfrom the modified Goldman equation. In the third and final phase,loss of K⁺ Na⁺ Cl⁺,sugars, and amino acids began to increase,soon followed by loss of betacyanin. Tissues that had lost their betacyanin during anoxia were irreversiblyinjured, as shown by rapid uptake of Evans Blue and afailureto take up K⁺ , Na⁺ and Cl⁺ during re–aeration. In contrast,tissues which had retained their betacyanin did not take upEvansBlue, but took up substantial amounts of K⁺ , Na⁺ , and Cl^–after re–aeration. After return to air for 1.5 h, tissuethat hadretained its betacyanin had a membrane potential of– 154 mV. Key words: Anoxia, beetroot, solute, membrane potential     

Growth and Osmoregulation of Chaetoceros muelleri in Relation to Salinity

The growth and osmoregulation of Chaetoceros muelleri Lemmermann(Bacillariophyceae) were investigated as a function of salinity.This centric diatom grew well over a wide range of salinityand required concentrations of NaCl above 10 mM for growth.Using gas chromatography- mass spectroscopy (GC-MS) analysisof cell extracts, we demonstrated that the alga contains anisomer of cyclohexanetetrol. The level of this isomer increasedwith increasing salinity. Levels of free amino acids also increasedwith increasing salinity, and quantitative determination withan amino acid analyzer revealed that the level of glutamic acidincreased the most with increases in salinity. Levels of intracellularK⁺ and Cl^– also increased significantly with increasesin salinity. Thus, in C. muelleri, not only organic solutessuch as the cyclohexanetetrol isomer and glutamic acid, butalso inorganic solutes such as K⁺ and Cl^– contribute toosmoregulation. (Received November 7, 1994; Accepted April 10, 1995)     

Effect of Salinity on Seed Germination,Growth, and Ion Accumulation of Atriplex patula (Chenopodiaceae)

Seeds and seedlings of the halophyte Atriplex patula were exposed to 0–2% NaCl to determine the effect of salt stress on germination and growth. Seeds germinated and plants survived and grew in solutions of up to 2.0% NaCl. Both seed germination and dry mass production were negatively affected by increased salinity. Dry mass production declined to 1% of controls and seed germination to 17% of controls in the 2% NaCl treatments, indicating that seeds were less inhibited than growing plants. Also, recovery treatments indicated that high salinity did not permanently injure seeds. Percent ash, and Na⁺ and Cl⁻ ions increased in shoots with each salt increment, while the K⁺ ion content decreased sharply. Atriplex patula is a facultative halophyte, and is limited to low and moderately saline sites because both seed germination and growth are severely reduced at salinities > 1% NaCl.     

Rate of Uptake of Potassium by Three Crop Species in Relation to Growth

Barley, ryegrass, and fodder radish were grown in flowing nutrientsolutions at four potassium concentrations, [K_e⁺], from 0.05to 4 mg I^–1. During the first 2 weeks after germinationthe response to [K_e⁺] (fodder radish > barley > ryegrass)depended on the potential relative growth rate, the ratio ofroot surface area to plant weight, and on the K⁺ flux into theroots. Subsequently, there was no effect of [K_e⁺] on growthrate within the range tested. The K⁺ flux decreased from 4–23? 10^–12 mol cm^–2 s^–1 in the first 2 weeksafter germination, when it was concentration-dependent, to 2–5? 10^–12 mol cm^–2 s^–1 after 4–5 weeks,when it became independent of [K_e⁺] down to 0.05 mg 1^–1.The results explain the importance of high [K_e⁺] and rapid rootgrowth during the first 2 weeks after seed germination.