Effect of high external NaCl concentration on ion transport within the shoot of Lupinus albus. II. Ions in phloem sap

Abstract. Phloem sap was collected from petioles of growing and fully expanded leaves of lupins exposed to 0–150 mol m⁻³ [NaCl]_ext, for various periods of time. Sap bled from growing leaves only after the turgor of the shoot was raised by applying pneumatic pressure to the root. Increased pressure was also needed to obtain sap from fully expanded leaves of plants at high [NaCl]_ext. Exposure to NaCl caused a rapid rise in the Na⁺ concentration in phloem sap to high levels. The Na⁺ concentration reached 20 mol m⁻³ within a day of exposure and reached a plateau of about 60 mol m⁻³ in plants at 50–150 mol m⁻³ [NaCl]_ext, after a week. There was a slower, smaller increase in the Cl⁻ concentration. K⁺ concentrations in phloem sap were not affected by [NaCl]_ext. Cl⁻ concentrations in phloem sap collected from growing leaves were similar to those from old leaves while Na⁺ concentrations were somewhat increased, suggesting that there was no reduction in the salt content of the phloem sap while it flowed within the shoot to the apex. Calculations of ion fluxes in xylem and phloem sap indicated that Na⁺ and Cl⁻ fluxes in the phloem from leaves of plants at high NaCl could be equal to those in the xylem. This prediction was borne out by observations that Na⁺ and Cl⁻ concentrations in recently expanded leaves remained constant.     

Endogenous abscisic acid levels are linked to decreased growth of bush bean plants treated with NaCl

This paper studies the relative importance of endogenous ABA and ion toxicity in the leaf growth inhibition caused by NaCl in salt-adapted and unadapted bush beans. Adaptation to salt-stress was achieved by germination of seeds in 75 m M NaCl, while unadapted plants were germinated in tap water. The adaptation process caused a transitory increase in leaf ABA, Na⁺ and Cl⁻ concentrations, while leaf expansion was inhibited. However, when grown for 8 or 13 days in 75 m M NaCl-containing nutrient solution, primary and first trifoliolate leaves of salt-adapted plants had greater areas than those of unadapted plants. Concentrations of ABA, Na⁺ and Cl⁻ in these leaves were lower in adapted plants, and a strong negative correlation between leaf expansion growth and either leaf Na⁺, Cl⁻ or ABA concentrations could be established. However, in the second trifoliolate leaves only the ABA, but not the Na⁺ or Cl⁻, concentrations were significantly correlated with leaf expansion. Our results suggest that salt-induced inhibition of leaf expansion growth in bush beans is mediated by ABA rather than Na⁺ or Cl⁻ toxicity. Moreover, the increase of ABA, induced by the salt-pretreatment, seems to play an important role in limiting the accumulation of Na⁺ and Cl⁻ in the leaves, leading to adaptation of bush beans to salt-stress.     

Sodium partitioning within the shoot of soybean

Uptake and partitioning of Na⁺ and Cl⁻ in plants of soybean ( Glycine max L. Merr. cv. Hodgson) exposed to moderate NaCl concentrations were studied over an 8-day period. Plants showed marked retention of Na⁺ in the stems and low transport to laminae of young leaves. The xylem sap ascending the main axis was progressively depleted in Na⁺. The oldest leaf greatly contributed to Na⁺ depletion of the sap flowing to younger leaves. These results in combination with estimates of phloem recirculation indicated that Na⁺ accumulation in the young leaf was prevented both by depletion of Na⁺ from the xylem stream, and by a high recirculation of Na⁺ via the phloem. However, this protection of young leaves was effective only for very mild salinity treatment.     

Response of Suaeda aegyptiaca to KCl, NaCl and Na2SO4 treatments

The response of Suaeda aegyptiaca (Hasselq.) Zoh. to various salinity treatments was tested in sand culture. Growth was promoted by NaCl and by Na₂SO₄ at all tested concentrations, but not by KCl. The effect of NaCl on growth was stronger than that of Na₂SO₄ and it increased gradually up to a 125 eq. m⁻³ optimum. Ion uptake was also affected by the different salts. Cl⁻ was taken up in similar quantities from KCl and from NaCl solutions and the content of the respective cations was also similar to one another. The presence of Na⁺ in the medium lowered the content of K⁺ in the plants and at the same time increased growth by as much as 900%. Transpiration was reduced and water use efficiency increased by Na⁺-salts. Highest water use efficiency was exhibited by plants which were treated with 125 eq. m⁻³ NaCl. It is concluded that Na⁺ at the macronutrient level has a specific promotive effect on the physiological processes of S. aegyptiaca. This effect is not due to replacement of K⁺ by Na⁺; neither can it be achieved by increasing the K⁺ concentration. Cl⁻ has an additional positive effect on growth of S. aegyptiaca. This effect is only expressed in the presence of Na⁺.     

Stomatal limitation of photosynthesis and reduced growth of the halophyte, Plantago maritima L., at high salinity

Abstract. Plantago maritima L. was grown at three levels of salinity, 50, 200, 350 mol m⁻³ NaCl, and the effects on growth, ion content and photosynthetic capacity were studied. Shoot and root dry weight, leaf production and leaf length were all substantially reduced in plants grown at high salinity. Total leaf area of plants grown at 350 mol m⁻³ NaCl was only 20% of that in plants at low salinity. Both the Na⁺ and K⁺ content of leaves and roots increased with external salinity. There was no change in the Na⁺/K⁺ ratio of leaves or roots at different salinity levels. Despite the large reductions in growth and high accumulation of Na⁺ ions, leaf photosynthetic rate was only slightly reduced by salinity stress. The reduction in photosynthesis was not caused by reduced biochemical capacity as judged by photosynthetic response to intercellular CO₂ and by ribulose-1,5-bisphosphate carboxylase activity, but was due to reduced leaf conductance and low intercellular CO₂ concentration. The increased stomatal limitation of photosynthesis resulted in higher water-use efficiency of plants grown at high salinity.     

Interactions between growth, uptake of Cl− and Na+, and water relations of plants in saline environments. II. Highly vacuolated cells

Abstract. In a highly saline environment high rates of ion uptake are required to generate sufficient osmotic pressure to maintain the turgor that is needed for the continued growth of plants. We estimate the rates of net uptake of Cl⁻ and Na⁺ required by growing cells to sustain cell expansion at an external NaCl concentration of 500 mol m⁻³. We also estimate the ion fluxes required to regulate turgor of expanding and fully expanded cells during diurnal changes in transpiration. Passive fluxes could contribute significantly to osmotic regulation, but active fluxes are still essential and would consume a substantial amount of energy. We discuss whether a limitation to growth at high salinity would arise from lack of energy, or from insufficient capacity for ion uptake. There is insufficient evidence to choose between these possibilities.     

Effects of chloride and sodium on gas exchange parameters and water relations of Citrus plants

Gas exchange parameters, water relations and Na⁺/Cl^- content were measured on leaves of one-year-old sweet orange ( Citrus sinensis [L.] Osbeck cv. Hamlin) seedlings grown at increasing levels of salinity. Different salts (NaCl, KCl and NaNO₃) were used to separate the effects of Cl⁻ and Na⁺ on the investigated parameters. The chloride salts reduced plant dry weight and increased defoliation. Accumulation of Cl⁻ in the leaf tissue caused a sharp reduction in photosynthesis and stomatal conductance. By contrast, these parameters were not affected by leaf Na⁺ concentrations of up to 478 m M in the tissue water. Leaf water potentials reached values near −1.8 MPa at high NaCl and KCl supplies. This reduction was offset by a decrease in the osmotic potential so that turgor was maintained at or above control values. The changes in osmotic potential were closely correlated with changes in leaf proline concentrations. Addition of Ca²⁺ (as calcium acetate) increased growth and halved defoliation of salt stressed plants. Furthermore, calcium acetate decreased the concentration of Cl⁻ and Na⁺ in the leaves, and increased photosynthesis and stomatal conductance. Calcium acetate also counteracted the reductions in leaf water and osmotic potentials induced by salinity. In addition, calcium acetate inhibited the accumulation of proline in the leaves which affected the reduction in osmotic potential. These results indicate that adverse effects of salinity in Citrus leaves are caused by accumulation of chloride.     

Changes in sodium and potassium in Nitellopsis cells treated with transient salt stress

Abstract. Nitellopsis cells grown in fresh water have a relatively low cytoplasmic Na⁺ (11 mol m⁻³) and high cytoplasmic K⁺ (90 mol m⁻³) content. A 30-min treatment with 100 mol m⁻³ external NaCl resulted in a high [Na⁺]_c (90 mol m⁻³) and a low [K⁺]_c (33 mol m⁻³), Subsequent addition of external Ca²⁺ (10 mol m⁻³) prevented Na⁺ influx and then [Na⁺]_c decreased slowly. Changes in [K⁺]_c were opposite to [Na⁺]_c. During the recovery time vacuolar Na⁺ increased, while vacuolar K⁺ decreased. Since all these processes proceeded also under ice-cold conditions, the restoration of original cytoplasmic ion compositions is suggested to be a passive nature. The notion that the passive movement of ions across the tonoplast can act as an effective and economic mechanism of salt tolerance under transient or under mild salt stress conditions is discussed.     

Salinity tolerance of Kosteletzkya virginica. I. Shoot growth, ion and water relations

Abstract. Kosteletzkya virginica (L.) Presl., a dicotyledonous halophyte native to brackish tidal marshes, was grown on nutrient solution containing 0. 85, 170 or 255 mol m^-3 NaCl, and the effects of external salinity on shoot growth and ion content of individual leaves were studied in successive harvests. Growth was stimulated by 85 mol m^-3 NaCl and was progressively reduced at the two higher salinities. Growth suppression at high salinity resulted principally from decreased leaf production and area, not from accelerated leaf death. As is characteristic of halophytic dicots. K. virginica accumulated inorganic ions in its leaves, particularly Na⁺ and K⁺. However, the Na⁺ concentration of individual leaves did not increase with time, but remained constant or even declined, seeming to be well-coordinated with changes in water content. A striking feature of the ion composition of salinized plants was the development of a dramatic gradient in sodium content, with Na⁺ partitioned away from the most actively growing leaves. Salt-treated plants exhibited a strong potassium affinity, with foliar K⁺ levels higher in salinized plants than unsalinized plants after an initial decrease. These results suggest that selective uptake and transport, foliar compartmentation of Na⁺ and K⁺ in opposite directions along the shoot axis, and the regulation of leaf salt loads over time to prevent build-up of toxic concentrations are whole-plant features which enable K. virginica to establish favourable K⁺-Na⁺ relations under saline conditions.     

Functional anatomy controls ion distribution in banana leaves: significance of Na+ seclusion at the leaf margins

Typical salt stress symptoms appear in banana ( Musa sp., cv. 'Grand Nain' AAA) only along the leaf margins. Mineral analysis of the dry matter of plants treated with increasing concentrations of KCl or NaCl revealed significant accumulation of Na⁺, but not of K⁺ or Cl^-, in the affected leaf margins. The differential distribution of the three ions suggests that water and ion movement out of the xylem is mostly symplastic and, in contrast to K⁺ and Cl^-, there exists considerable resistance to the flow of Na⁺ from the xylem to the adjacent mesophyll and epidermis. The parallel veins of the lamina are enclosed by several layers of bundle sheath parenchyma; in contrast, the large vascular bundle that encircles the entire lamina, and into which the parallel veins merge, lacks a complete bundle sheath. Xylem sap containing a high concentration of Na⁺ is 'pulled' by water tension from the marginal vein back into the adjacent mesophyll without having to cross a layer of parenchyma tissue. When the marginal vein was dissected from the lamina, the pattern of Na⁺ distribution in the margins changed markedly. The distinct anatomy of the marginal vein plays a major role in the accumulation of Na⁺ in the margins, with the latter serving as a 'dumping site' for toxic molecules.     

Salinity response of a freshwater charophyte, Chara vulgaris

Abstract. Chara vulgaris L. growing in an oligohaline lake was adapted to laboratory conditions and subjected to long-term salinity treatments ranging from 0 to 350 mol m ³ NaCl added to the lake water (40–680 mosmol kg ¹). Osmotic potential and concentration of the main osmotically active solutes (K⁺, Na⁺, Mg²⁺, Cl and sucrose) in the vacuolar sap of the central internodal cells were estimated. C. vulgaris did regulate turgor but incompletely. Turgor decreased from 335 mosmol kg ¹ under control conditions to 52–111 mosmol kg ¹ at 350 mol m ³ NaCl. The enhancement of πⁱ was achieved by increase in both ions and sucrose. Sterile and fertile plants differed in their response to osmotic stress. In sterile plants, the ions accounted for about 87% of the vacuolar osmotic potential. The increase of πⁱ under osmotic stress was exclusively due to an accumulation of Na⁺ and Cl^-. In fertile plants, sucrose accounted for about 35% of πⁱ and ions for about 51% Under osmotic stress, sucrose content increased together with the ionic content of Na⁺ and Cl^-.     

Role of sodium in the ABA‐mediated long‐term growth response of bean to salt stress

Long‐term salt effects on plant growth have often been related to direct ion toxicity due to the accumulation of high ion concentrations in plant tissue. This work examines the relative importance of endogenous ABA, as well as Na⁺ and Cl⁻ toxicity, in the inhibition of leaf growth and photosynthesis, in bean plants grown at 1, 25, 50 and 75 m M NaCl until the fruit‐bearing stage. All salt‐treated plants showed very high leaf Cl⁻ concentrations, with little difference between plants exposed to 50 or 75 m M NaCl. The 25 and 50 mM salt‐treated plants were able to successfully exclude Na⁺ from their leaves, and only suffered an initial decline in the rate of leaf growth. Plants exposed to 75 m M NaCl showed an increase in Na⁺ leaf concentrations with an accompanying decrease in growth and photosynthesis as salt exposure progressed. A high correlation was found between leaf Na⁺ and leaf growth. Leaf ABA significantly increased with salt supply, and was highly correlated with both leaf Na⁺ and leaf growth. Our results suggest that in bean plants under long‐term salt stress, leaf ABA may participate in the regulation of leaf growth, and leaf Na⁺ would be at least partly responsible for increased ABA levels.     

Physiological function of water channels as affected by salinity in roots of paprika pepper

The sap flow (Jv) and the osmotic pressure-dependent hydraulic conductance (L₀) of detached exuding root systems from paprika pepper plants (cv. Albar) were measured. Plants stressed with NaCl (30 m M ) and with six times the macronutrients of the Hoagland nutrient solution (6×HNS) were compared with controls grown in complete Hoagland nutrient solution. Jv of +NaCl and +6×HNS plants decreased markedly, but recovered to values similar to those of controls after removal of the treatments. Hydraulic conductance L₀ was always less in NaCl plants than in controls and 6×HNS. A total increase in the ion concentration of the xylem (except Na⁺ and Cl⁻) was observed with both treatments. In control and 6×HNS plants, HgCl₂ treatment (50 μ M ) caused a sharp decline in L₀ to values similar to those of NaCl-stressed roots, but were restored by treating with 5 m M dithiothreitol (DTT). However, in NaCl roots only a slight effect of Hg²⁺ and DTT was observed. In each treatment, there was no difference in the flux of K⁺ into the xylem after HgCl₂ and DTT application. The results suggest that NaCl decreased L₀ of the roots by reducing either the activity or abundance of Hg-sensitive water channels. The putative reduction in water-channel function of NaCl-treated plants did not seem to be due to the osmotic effect.     

Physiological effects of Na2SO4 and NaCl on callus cultures of Brassica campestris (Chinese cabbage)

Hypocotyl-derived callus cultures of Brassica campestris L. ssp. pekinensis cv. Kim-jung (Chinese cabbage) were grown on Murashige and Skoog medium containing no additional salt, NaCl or Na₂SO₄. Na₂SO₄ was more than twice as inhibitory in comparison to the same concentration of NaCl when growth and fresh:dry weight ratios of established callus were measured. Levels of protein, starch, sucrose and α-amino nitrogen were not significantly altered in salt-grown callus. Concentrations of reducing sugars and chlorophyll were 2–3 times greater in callus grown on either salt. Proline concentration increased 15–20 fold on the highest levels of salt. Final concentrations (reached in 20–24 days) were closely correlated to the initial Na⁺ concentration of the medium, regardless of salt type. The osmotic potential in callus transferred to NaCl or Na₂SO₄ reached a maximum negative value after 16 days. For both salts, subsequent increases were correlated to increases in fresh:dry weight and growth. On both salts, turgor remained relatively constant (0. 6–0.75 MPa). Changes in Na⁺, K⁺, Mg²⁺ and Ca²⁺ content were correlated to initial Na⁺ concentration in the medium, not salt type. Accumulation of Na⁺ was accompanied by loss of K⁺ and Mg²⁺. Six to seven times less sulfate was measured in callus grown on Na₂SO₄ than chloride in callus grown on similar concentrations of NaCl.     

Do leaves contribute to the abscisic acid present in the xylem sap of ‘draughted’ sunflower plants?

Abstract. The root systems of 30-d-old sunflower plants were treated with polyethylene glycol (PEG; osmotic potential - 1.0MPa) for 2h, causing mild and transient wilting. Ten minutes before this treatment was applied, half the plants were defoliated. At varying times after the imposition of the PEG 'drought stimulus', the plant stems were cut and the sap exudate was collected and analysed for abscisic acid (ABA), using an elisa method. When stems were cut 2.25h after the treatments were applied, the ABA concentration in the sap of the controls did not vary with time: the mean concentration was 10.7 ± 1.0μ, mol m⁻³. However, in the treated plants, the first sample contained 78.1 ± 10.1 μmol m⁻³, decreasing to 13.6 ± 2.8 μmol m⁻³ over 8.75h. Defoliation did not affect the ABA concentration in the sap. When stems were cut at varying times (up to 25h) after treatment, the PEG treatment again caused an immediate increase in the ABA concentration in the sap, from 20 ± 1 to 136 ± 21 μmol m⁻³. However, defoliation reduced this increase, but only in plants sampled 4–25h after treatment. We conclude that, after the PEG treatment to the roots, the initial increase of the ABA content of sap, and its attenuation with time, may be ascribed to synthesis in the roots whereas, thereafter, ABA derived from the leaves makes a major contribution to the ABA found in the xylem sap.     

Excretion of ions (Cd2+, Li+, Na+ and Cl−) by Tamarix aphylla

Excretion of minerals by the NaCl-resistant and comparatively cadmium-resistant tree Tamarix aphylla (L.) Karst, was investigated. Cd²⁺ was excreted by plants exposed for 1–10 days to 9 or 45 μ M Cd²⁺ solutions. Excretion of this toxic ion increased considerably with time but was less than 5% of the quantities that had been accumulated in the shoots. Excretion of Na⁺ and Cl⁻ was positively correlated with NaCl concentration (1.5, 10, 50 m M ) of the medium. The Na⁺/Cl⁻ ratios of the excrete were positively correlated with the concentration of the treatment solution. Ca²⁺ excretion decreased with increasing NaCl concentrations of the solution. Excretion of K⁺ and Mg²⁺ was only little affected by NaCl. Excretion of Li⁺ occurred whenever this element was supplied in the uptake solution; daily excretion rates of Li⁺ increased with time. The ecological significance of excretion is discussed in relation to the low selectivity of the mechanism in T. aphylla .     

Enhancement of salt tolerance in soybean with NaCl pretreatment

Acclimation response to salt stress in soybean ( Glycine max [L.] Merr. cv. Lee) was found in this study. Soybean seedlings were exposed to 0, 34 and 68 m M NaCl for 23 days (pretreatment), thereafter plants were exposed to 0, 68 and 137 m M NaCl until maturity (main treatment). There was no effect of pretreatment on growth, but at 137 m M NaCl, Na⁺ concentration in leaves of the plants pretreated with 34 m M NaCl was lower than that of plants pretreated with 0 and 68 m M NaCl. Furthermore, the survival rate under 137 m M NaCl improved with the 34 m M NaCl pretreatment. Therefore, it is possible that soybean can acclimate to salt stress by its increased survival rate, without showing any improvement in growth. The regulation of Na⁺ or Cl⁻ concentration in leaves could be one of the possible factors involved in salt acclimation of soybean.     

Influence of exogenously supplied potassium and sodium salts on the abscisic acid-induced proline accumulation in barley leaf segments

A stimulation of the abscisic acid (ABA)-induced increase in proline was observed in leaf segments of barley ( Hordeum vulgare L. cv. Georgie) if K⁺ or Na⁺ were supplied in the external medium as salts of monovalent anions such as NO₃, Br, Cr and I, but not when sulphate or phosphate were used. To a lesser extent, the effect was evident also with RbCl, but it did not occur when chlorides of Li⁺. Cs⁺, NH₄⁺, Mg:⁺ and Ca²⁺ were used. Both KC1 and NaCl in the concentration range 2–100 m M influence the ABA-dependent proline accumulation to the same extent; the increase induced was about 100% at 10 m M , and reached a maximum between 60 and 100 m M. The effect is not due to the osmotic activity of the salts and does not seem to depend on changes in K⁺ and Na⁺ levels within the leaf tissue, but it is somehow linked to their external concentration. The existence of a specific interaction between ABA and K⁺ or Na⁺, possibly at the cell membrane level, is proposed.     

Salt stress in cultured rice cells: effects of proline and abscisic acid

Abstract. The presence of 1 and 10 mol m⁻³ proline in media containing 100 and 200 mol m⁻³ of NaCl, had little effect on the growth of salt-adapted callus of rice. However, in such callus proline accumulation was stimulated by 10 mol m⁻³ proline in the presence of 100 mol m⁻³ NaCl. On the other hand, with 100 mol m⁻³ NaCl, both 1 and 10 mol m⁻³ proline significantly increased both the growth and proline content of salt-unadapted callus. On replacing NaCl with KCl (100 and 200 mol m⁻³), growth of saltadapted as well as unadapted callus was inhibited, but the presence of 10 mol m⁻³ proline had an ameliorating effect. Abscisic acid (ABA) supressed the growth of both salt-adapted and unadapted callus of rice in the absence of salt stress. ABA inhibited the growth of callus adapted to and grown in 100 and 200 mol m⁻³ of NaCl or when it was replaced by equimolar concentrations of KCl. Growth of 100 mol m⁻³ NaCl adapted cells was inhibited when they were transferred to a medium containing 200 mol m⁻³ of NaCl, but in the presence of ABA it was stimulated. ABA increased the growth of unadapted cells when subjected to different salts. Also, ABA accelerated the adaptation of cells exposed to salt but not to water deficits imposed by nonionic solutes.     

Impairment of Na+,K+-ATPase activity following enterotoxigenic Campylobacter jejuni infection: changes in Na+, Cl− and 3-O-methyl-D-glucose transport in vitro, in rat ileum

Abstract Unidirectional fluxes of Na⁺, Cl⁻ and 3-O-methyl-D-glucose (3-MG) were measured in vitro across Campylobacter jejuni live culture-infected and control rat ileal short-circuited tissues by the Using Chamber technique. Net secretion of Na⁺ and enhanced secretion of Cl⁻ ions was observed in the infected animals ( P < 0.001, n =6) as compared to the net absorption of Na⁺ and marginal secretion of Cl⁻ ions in the control animals. There was a significant decrease in the mucosal-to-serosal fluxes of 3-MG in C. jejuni -infected rat ileum. The specific Na⁺,K⁺-ATPase activity when measured biochemically in the membrane-rich fraction of enterocytes was found to be significantly lower (58%) in the infected group as compared to the control group ( P < 0.001). Our results therefore suggest that infection with an enterotoxigenic C. jejuni inhibits the Na⁺,K⁺-ATPase activity in rat enterocytes. The impairment of Na⁺,K⁺-ATPase activity thus appears to induce a secondary change in Na⁺,Cl⁻ and 3-MG transport in vitro in rat ileum.