Effect of NaCl salinity on growth and mineral partitioning in Quercus robur L., a rhythmically growing species

 Sodium salt sensitivity of common oak (Quercus robur L.) was evaluated in hydroponic culture using INRA-Morizet solution. Addition of NaCl to the nutrient solution reduced only length and weight of roots and first flush stems. In contrast, the second flush was properly expanded even in the presence of 40 mM of NaCl in culture medium. Both leaf number and leaf area were not affected by increasing salt concentration in medium culture while this increase induced significant leaf damage especially in first flush leaves. Stem starch storage was reduced only at 40 mM NaCl treatment. Common oak seedlings seemed to be able to better compartmentalize sodium than chloride when the NaCl concentration increased in the medium culture. Chloride presented a lower uptake than sodium. Sodium was preferentially accumulated in roots and this accumulation occurred at the expense of potassium uptake. The decrease of ATP content in leaves of common oak seedlings submitted to NaCl treatments could indicate that it was used for sodium exclusion out of the leaves, especially in second flush leaves. Relationships between growth responses, starch and mineral element distribution in common oak seedlings will be discussed. Received: 20 November 1997 / Accepted: 3 March 1998     

Functional Aspects of the Salt Glands of the Plumbaginaceae

X-ray microanalysis and diffuse reflectance infrared Fouriertransform spectrometry were used to determine the presence andratios of elements in salt secretions from salt glands of greenhouseand experimentally-manipulated leaves of five species of thePlumbaginaceae Sodium, magnesium, silica, sulphur, phosphorus,chloride, potassium, calcium and carbonate were detected insecretions of greenhouse-grown plants. The salt glands of excisedleaves challenged by solutions of KI, KCl, NaCl, Na₂SO₄, MgCl₂and MgSO₄ secreted principally the ions of the challenging solutions. Key words: Ion transport, secretion     

Salinity tolerance of citrus rootstocks: Effects of salt on root and leaf mineral concentrations

The effects of three concentrations of sodium chloride (NaCl) on seven citrus rootstocks were studied under greenhouse conditions. Leaf and root mineral concentrations and seedling growth were measured. Sodium chloride was added to the nutrient solution to achieve final osmotic potentials of –0.10, –0.20, and –0.35 MPa. Increasing the concentration of NaCl in the nutrition solution reduced growth proportionally and altered leaf and root mineral concentrations of all rootstocks. Significant differences in leaf and root mineral concentration among rootstocks were also found under stressed and non-stressed conditions. Salinity caused the greatest growth reduction in Milam lemon and trifoliate orange and the least reduction in sour orange and Cleopatra mandarin. No specific nutrient deficiency was the sole factor reducing growth and causing injury to citrus rootstocks. Sodium chloride sensitivity of citrus rootstocks in terms of leaf burn symptoms and growth reduction could be attributed more to Cl than to Na. Sodium and Cl concentrations were greater in the leaves than in the roots, particularly at the medium and high salinity levels. Root Cl was not useful for assessing injury because no differences were found in root Cl concentrations among rootstocks. Increasing salinity level did not affect the level of N and Ca in the roots but did reduce N and Ca levels in the leaves. No relationship in mineral concentration or accumulation seemed to exist between citrus leaves and roots. At the –0.10 MPa salinity level, sour orange, rough lemon, and Milam were not able to exclude either Na or Cl from their leaves. Trifoliate orange and its two hybrids (Swingle citrumelo and Carrizo citrange) excluded Na at the lowest salt level used, but were unable to exclude Na at the higher salinity levels. Similarly, Cleopatra mandarin excluded Cl at the lowest salt level, but was not able to exclude Cl at higher salt concentrations. Hence, the ability of citrus rootstocks to exclude Na or Cl breaks down at higher salt concentrations.Florida Agricultural Experiment Station Journal Series No. R-02276.     

Salt excretion from the salt glands in Rhodes grass (Chloris gayana Kunth) as evidenced by low-vacuum scanning electron microscopy

Some salt-tolerant plants belonging to the Poaceae excrete salts to the leaf surfaces under salinity conditions, and the bicellular glands on their leaf surfaces have been postulated to excrete salt. However, clear evidence of the salt excretion from these bicellular salt glands has not been shown at the electron-microscope level because soluble attachments on the leaf surface are completely removed during specimen preparation for conventional electron microscopy. To determine whether the bicellular salt glands actually excrete salt, we examined the leaves of Rhodes grass (Chloris gayana Kunth), Poaceae, by scanning electron microscopy in a low-vacuum mode, which allows to observe specimens without preparation procedures. Unwashed and washed fresh leaf surfaces were examined, and excreted materials on the leaf surface were analyzed by energy dispersive X-ray spectrometry. On the unwashed leaf surfaces, globular materials were observed arranged along the same lines as the macrohairs of the leaf surface, but the salt glands were hardly observed. After leaf surfaces were washed, the globular materials disappeared, and the salt glands appeared localized at the same lines as the macrohairs. Density of the globular materials observed under unwashed conditions and the salt glands under washed conditions was equal. These findings indicate that the glands indeed excrete globular materials just above their cap cells. The excreted materials contained sodium, chlorine, and potassium, and the counts of sodium and chlorine was increased greatly with NaCl treatment of the plants. After removing the excreted materials, most of the cap cells of the salt glands were smooth globular, without ruptures in their cuticle. We conclude that the leaves of Rhodes grass indeed excrete salt from the bicellular salt glands, but without rupturing the cuticle on the cap cell.     

Mechanism of salt tolerance in wild rice (Oryza coarctata Roxb)

Summary Oryza coarctata, a highly salt-resistant wild rice species, is commonly found on the banks of coastal rivers in India. This species can also withstand saline water (20 to 40 dSm⁻¹ E.C) submergence for quite a long period. It was revealed thatO. coarctata has some special unicellular salt hairs (trichomes) on the adaxial surface of the leaves, by which they efficiently maintain a low concentration of toxic salts in the plant tissue. Sodium and chloride were the dominant ions in the excreted material but they also excrete potassium, magnesium and calcium. With the increase in soil salinity sodium, magnesium and chloride excretion increased.O. coarctata maintained the optimum mineral concentration in its tissues. Maximum accumulation of potassium was observed in the leaves. With the increase in salt stress total biomass production and osmotic potential increased over control but there was no change in the moisture percentage of leaves.     

盐胁迫对獐茅生长及Na+和K+含量的影响

用含0～200mmol/L浓度的NaCl的Hoagland培养液处理獐茅幼苗,处理17 d后测定一些生理指标,结果表明獐茅的生长受NaCl抑制程度随浓度增加而增大,没观察到最适盐浓度,且叶片较根部对盐分更敏感;有机干重的比例增大表明有机物在渗透调节中的贡献随之增大.植株在加喷对盐腺向体外排盐有抑制作用的溶液-苯硫酸胆碱100mmol/L后,Na 、K 含量在叶片内增加,而在分泌物中的量降低,并表现出生长进一步受到抑制.X-ray微区分析结果表明獐茅可以将Na 区域化到盐腺细胞,以便将其分泌到体外.     

Intrinsic water use efficiency controls the adaptation to high salinity in a semi-arid adapted plant,henna (Lawsonia inermis L.)

Adaptation to salinity of a semi-arid inhabitant plant, henna, is studied. The salt tolerance mechanisms are evaluated in the belief that gas exchange (water vapor and CO₂) should play a key role on its adaptation to salt stress because of the strong evaporation conditions and soil water deficit in its natural area of distribution. We grow henna plants hydroponically under controlled climate conditions and expose them to control (0 mM NaCl), and two levels of salinity; medium (75 mM NaCl) and high (150 mM NaCl). Relative growth rate (RGR), biomass production, whole plant and leaf structure and ultrastructure adaptation, gas exchange, chlorophyll fluorescence, nutrients location in leaf tissue and its balance in the plant are studied. RGR and total biomass decreased as NaCl concentration increased in the nutrient solution. At 75 mM NaCl root biomass was not affected by salinity and RGR reached similar values to control plants at the end of the experiment. At this salinity level henna plant responded to salinity decreasing shoot to root ratio, increasing leaf specific mass (LSM) and intrinsic water use efficiency (iWUE), and accumulating high concentrations of Na⁺ and Cl⁻ in leaves and root. At 150 mM NaCl growth was severely reduced but plants reached the reproductive phase. At this salinity level, no further decrease in shoot to root ratio or increase in LSM was observed, but plants increased iWUE, maintaining water status and leaf and root Na⁺ and Cl⁻ concentrations were lower than expected. Moreover, plants at 150 mM NaCl reallocated carbon to the root at the expense of the shoot. The effective PSII quantum yield [Y(II)] and the quantum yield of non-regulated energy dissipation [Y(NO)] were recovered over time of exposure to salinity. Overall, iWUE seems to be determinant in the adaptation of henna plant to high salinity level, when morphological adaptation fails.     

Ultrastructure of Aeluropus littoralis leaf salt glands under NaCl stress

The effects of salt uptake on the morphology and ultrastructure of leaf salt glands were investigated in Aeluropus littoralis plants grown for two months in the presence of 400 mM NaCl. The salt gland is composed of two linked cells, as observed in some other studied Poaceae species. The cap cell, which protrudes from the leaf surface, is smaller than the basal cell, which is embedded in the leaf mesophyll tissues and bears the former. The cuticle over the cap cell is frequently separated from the cell wall to form a cavity where salts accumulate prior to excretion. The basal cell cytoplasm contains an extensive intricate or partitioning membrane system that is probably involved in the excretion process, which is absent from the cap cell. The intricate membrane system seems to be elongated and heavily loaded with salt. The presence of 400 mM NaCl induced the disappearance of the collecting chamber over the glands and an increase in the number of vacuoles and their size in both gland cells. In the basal cell, salt greatly increased both the density and size of the intricate membrane system. The electron density of both gland cells observed under salt treatment reflects a high activity. All these changes probably constitute special adaptations for dealing with salt accumulation in the leaves. Despite the high salt concentration used, no serious damage occurred in A. littoralis salt gland ultrastructure, which consolidates the assumption that they are naturally designated for this purpose.     

Growth and mineral content of coriander (<Emphasis Type="Italic">Coriandrum sativum</Emphasis> L.) plants under mild salinity with different salts

Soil salinity is a complex issue in which various anions and cations contribute to have a general adverse effect on plant growth. In the present study, effects of salinity from various salts including sodium chloride (NaCl), potassium chloride?+?sodium chloride?+?calcium chloride (KCl?+?NaCl?+?CaCl₂), potassium sulfate?+?magnesium nitrate (K₂SO₄?+?Mg(NO₃)₂) at two electric conductivities (EC) of 2 and 4 dS m^?1 of irrigation water, and a distilled water control were evaluated on coriander plants (Coriandrum sativum L.). At EC?=?2, all salts increased plant yield (shoot fresh weight) than control. Most growth traits including plant height, shoot fresh and dry weight, leaf SPAD value and vitamin C, leaf K, Mg and P concentrations were increased by K₂SO₄?+?MgNO₃, and remained unchanged by KCl?+?NaCl?+?CaCl₂ treatment (except reduced plant height). Leaf’s zinc concentration reduced by either treatment. Even sodium chloride at EC?=?2 showed some beneficial effects on leaf chlorophyll index, root fresh weight, leaf’s calcium and phosphorus concentration; however, most traits remained unchanged than control. Treatment of plants with NaCl or KCl?+?NaCl?+?CaCl₂ at either EC increased the number of flowered shoots and leaf proline content than control. Most growth and quality traits including leaf minerals and vitamin C content were reduced by NaCl at EC?=?4; however, shoot fresh and dry weights remained unchanged than control. Plant root fresh weight increased by NaCl at EC?=?2 and decreased at EC?=?4 than control. At EC?=?4, shoot dry weight was increased and leaf Ca, P, Zn and Mn were decreased by KCl?+?NaCl?+?CaCl₂, whereas shoot dry weight, leaf SPAD value and vitamin C content, leaf Mg and P were increased and leaf Zn was decreased by K₂SO₄?+?MgNO₃ than control. The results indicate that in contrast to sodium chloride, the salinity effects of other salts can not be detrimental on coriander plant growth.     

Salinity Stiffens the Epidermal Cell Walls of Salt-Stressed Maize Leaves: Is the Epidermis Growth-Restricting?

As a result of salt (NaCl)-stress, sensitive varieties of maize (Zea mays L.) respond with a strong inhibition of organ growth. The reduction of leaf elongation investigated here has several causes, including a modification of the mechanical properties of the cell wall. Among the various tissues that form the leaf, the epidermis plays a special role in controlling organ growth, because it is thought to form a rigid outer leaf coat that can restrict elongation by interacting with the inner cell layers. This study was designed to determine whether growth-related changes in the leaf epidermis and its cell wall correspond to the overall reduction in cell expansion of maize leaves during an osmotic stress-phase induced by salt treatment. Two different maize varieties contrasting in their degree of salt resistance (i.e., the hybrids Lector vs. SR03) were compared in order to identify physiological features contributing to resistance towards salinity. Wall loosening-related parameters, such as the capacity of the epidermal cell wall to expand, β-expansin abundance and apoplastic pH values, were analysed. Our data demonstrate that, in the salt-tolerant maize hybrid which maintained leaf growth under salinity, the epidermal cell wall was more extensible under salt stress. This was associated with a shift of the epidermal apoplastic pH into a range more favourable for acid growth. The more sensitive hybrid that displayed a pronounced leaf growth-reduction was shown to have stiffer epidermal cell walls under stress. This may be attributable to the reduced abundance of cell wall-loosening β-expansin proteins following a high salinity-treatment in the nutrient solution (100 mM NaCl, 8 days). This study clearly documents that salt stress impairs epidermal wall-loosening in growth-reduced maize leaves.     

Growth and Water Relations of Lotus Creticus Creticus Plants as Affected by Salinity

Young plants of Lotus creticus creticus growing in a hydroponic culture were submitted to 0, 70 and 140 mM NaCl treatments for 28 d and the growth and ecophysiological characteristics of these plants have been studied. The growth of Lotus plants was not affected by salinity when applied for a short period (about 15 d); however, 140 mM NaCl induced a decrease in shoot RGR at the end of the treatment. The root growth was not decreased, even it was stimulated by 140 mM NaCl. The osmotic adjustment of Lotus plants at 70 and 140 mM NaCl maintained constant pressure potential, avoiding the visual wilting. For a similar leaf water potential, cuticular transpiration of salinized plants was lower than in control plants due to the salinity effect on the cuticle. Moreover, the presence of hairy leaves (60 and 160 trichomes per mm² in young and adult leaves, respectively) allows keeping almost 81 % of sprayed water and absorbing the 9 % of the water retained, decreased the epidermal conductance to water vapour diffusion.     

Effects of salinity and benzyl adenine on development and function of microhairs of Zea mays L

Bicellular microhairs are present on the surfaces of leaves of grasses with the exception of the Pooideae. In some halophytic grasses, these glandular hairs secrete salt, suggesting the intriguing question ‘can the microhairs of grasses that do not normally encounter salinity also secrete salt?’ Microhairs were counted in replicas of the adaxial and abaxial surfaces of leaves of various ages of maize plants growing either in the absence of salt or in the presence of 40, 80 or 120 mM NaCl. The number of microhairs per unit area of adaxial leaf surface of the youngest leaf almost doubled as the salinity increased from zero to 120 mM NaCl; on the abaxial surface, the number of microhairs increased by 50%. Spraying this leaf with benzyl adenine (BA) caused, when averaged across salinities and surfaces, a 32% increase in the number of microhairs. Salinity reduced leaf area but in all the salinity treatments, spraying with BA increased the total number of microhairs per leaf. Washing leaves of plants provided estimates of the loss of salt from those leaves. There were large differences between the Na:K molar ratios in the washing solution and the leaf tissue, indicating a high selectivity for sodium over potassium for loss from the leaf. BA did not influence the efficiency of salt loss, expressed per microhair, at any salinity level, but did increase loss per leaf. Thus, BA increased salt loss from plants due to its influence on the number of microhairs and leaf area, but not due to its effect on the efficiency of the secretion process per se.     

Differential responses of antioxidative enzymes and lipid peroxidation to salt stress in salt-tolerant Plantago maritima and salt-sensitive Plantago media

The changes in plant growth, relative water content (RWC), stomatal conductance, lipid peroxidation and antioxidant system in relation to the tolerance to salt stress were investigated in salt-tolerant Plantago maritima and salt-sensitive Plantago media. The 60 days old P. maritima and P. media seedlings were subjected to 0, 100 and 200 mM NaCl for 7 days. Reduction in shoot length was higher in P. media than in P. maritima after exposure to 200 mM NaCl, but 100 mM NaCl treatment did not show any effect on shoot length of P. maritima. Shoot dry weight decreased in P. media and did not change in P. maritima. Two hundred millimolar NaCl treatment had no effect on leaf RWC in P. maritima, but it was reduced in P. media. Salt stress caused reduction in stomatal conductance being more pronounced in P. media than in P. maritima. Activities of superoxide dismutase (SOD; EC 1.15.1.1), catalase (CAT; EC 1.11.1.6), glutathione reductase (GR; EC 1.6.4.2) decreased in P. media with increasing salinity. Ascorbate peroxidase (APX; EC 1.11.1.11) activity in leaves of P. media was increased and showed no change under 100 and 200 mM NaCl, respectively. However, activities of CAT, APX and GR increased under 200 mM NaCl while their activities did not change under 100 mM NaCl in P. maritima. SOD activity in leaves of P. maritima increased with increasing salinity. Concomitant with this, four SOD activity bands were identified in leaves of P. maritima, two bands only were observed in P. media. Peroxidase (POX; EC 1.11.1.7) activity increased under both salt concentrations in P. maritima, but only under 200 mM NaCl in P. media. Confirming this, five POX activity bands were identified in leaves of P. maritima, but only two bands were determined in P. media. Malondialdehyde levels in the leaves increased under salt stress in P. media but showed no change and decreased in P. maritima at 100 and 200 mM NaCl, respectively. These results suggest that the salt-tolerant P. maritima showed a better protection mechanism against oxidative damage caused by salt stress by its higher induced activities of antioxidant enzymes than the salt-sensitive P. media.     

Growth responses of Stylosanthes humilis (Fabaceae) populations to saline stress

The growth responses to salinity of seedlings of six populations of Stylosanthes humilis from three ecogeographic regions of Northeast Brazil, characterized by wet or semi-arid climate, were analyzed following 28 days in solution cultures at concentrations of 0, 40, 80 and 120 mM NaCl. Root and shoot mass, shoot length and characters of foliar damage (number of leaves with chlorosis and necrosis) of the populations were more affected by increasing NaCl than numbers of leaves and branches and root length. Although S. humilis has been considered a salt sensitive legume, significant differences in salt tolerance between populations were found. The estimated concentrations which reduced shoot dry mass by 50 and 25% varied between populations from 84 to 108 and from 49 to 83 mM NaCl, respectively. Salt tolerance in S. humilis during the initial growth stage was lower than the reported one for germination. With one exception, populations from semi-arid climate with saline soils showed higher salt tolerance than those from non saline soils. The results suggest that salt tolerance in these populations is mainly associated with the occurrence of salinity in the soil of their provenance.     

盐胁迫下海马齿叶片结构变化

用石蜡切片法制片、光学显微镜观察了海马齿植物营养器官--叶片的盐适应结构变化,以明确盐生植物对盐渍生境适应的叶片结构变化特征,为盐生植物的耐盐机理研究提供依据.结果表明:(1)海马齿植物叶片表现出许多适应干旱和盐渍环境的特点,其基本特征为:叶片肉质化,为典型的等面叶;栅栏组织发达,且含有大量叶绿体;叶表皮气孔微下陷,叶表皮细胞外壁的角质层较薄,表皮细胞大小不等,外切向壁外凸,参差不齐,有些表皮细胞特化为泡状细胞,其数量与盐胁迫的浓度呈正相关.(2)叶的海绵组织中含有大量的薄壁细胞,幼叶海绵组织的薄壁细胞在0.5%～2.5% NaCl胁迫下均变大,且数量也增加;而老叶海绵组织的薄壁细胞只有在低浓度(0.5% NaCl)的盐胁迫下变大,而在高浓度下其薄壁细胞反而变小或成不规则形状.(3)盐晶广泛分布在海马齿的叶肉组织细胞内,且其数量随着盐胁迫浓度增加而增加.     

NaCl tolerance in <Emphasis Type="Italic">Lycopersicon pennellii</Emphasis> introgression lines: QTL related to physiological responses

The growth and ion content of salt sensitive Lycopersicon esculentum Mill. cv. M82 and salt tolerant L. pennellii Correll accession LA716 were examined under both control and stress conditions (150 mM NaCl). L. esculentum grew more vigorously than L. pennellii under optimal conditions, however, L. pennellii was able to maintain its growth better than cultivated tomato when the plants were exposed to salinity. Sodium content of both L. esculentum and L. pennellii increased as a result of NaCl stress. In addition, both species showed reduced potassium and calcium content due to salinity. The physiological traits were also measured in a population of 52 L. pennellii introgression lines grown under both normal and stress conditions. A total of 311 quantitative trait loci (QTL) were identified for the studied traits: plant height, stem diameter, leaf number, leaf and root fresh and dry mass, and sodium, potassium and calcium contents. Some of the loci (124) were identified under both control and stress conditions while 86 QTL were identified only under non-stress conditions and 101 loci were identified only under NaCl stress.     

THE EFFECT OF MODERATE SALT STRESS ON LEAF ANATOMY IN HIBISCUS CANNABINUS (KENAF) AND ITS RELATION TO LEAF AREA

Kenaf responded to salt stress in a manner that was typical of moderately salt tolerant non-halophytes. Increase in leaf area was more sensitive to salinity than either leaf emergence rate or dry matter accumulation. Dry weight was reduced only above a threshold of approximately 37 mM NaCl while leaf area was already significantly reduced at this salt concentration. Measurement of epidermal cell cross sectional area and epidermal cell numbers showed that the salt induced reduction in leaf area was due primarily to smaller epidermal cell size. Epidermal cell numbers were also significantly reduced by salinity. Stomatal density increased with increasing salt stress and there was no effect on leaf thickness.     

Abscisic Acid Reduces Leaf Abscission and Increases Salt Tolerance in Citrus Plants

This paper describes the physiological effects of abscisic acid (ABA) and 100 mM NaCl on citrus plants. Water potential, leaf abscission, ethylene production, photosynthetic rate, stomatal conductance, and chloride accumulation in roots and leaves were measured in plants of Salustiana scion [Citrus sinensis (L) Osbeck] grafted onto Carrizo citrange (Citrus sinensis [L.] Osbeck × Poncirus trifoliata [L.] Raf) rootstock. Plants under salt stress accumulated high amounts of chloride, increased ethylene production, and induced leaf abscission. Stomatal conductance and photosynthetic rates rapidly dropped after salinization. The addition of 10 mM ABA to the nutrient solution 10 days before the exposure to salt stress reduced ethylene release and leaf abscission. These effects were probably due to a decrease in the accumulation of toxic Cl- ions in leaves. In non-salinized plants, ABA reduced stomatal conductance and CO2 assimilation, whereas in salinized plants the treatment slightly increased these two parameters. The results suggest a protective role for ABA in citrus under salinity.     

Osmoregulation in Rhizobium meliloti: inhibition of growth by salts

A defined medium of low osmolarity was developed permitting growth of Rhizobium meliloti with generation times of approximately 2.8 h doubling^-1. The effects of sodium, potassium, magnesium, ammonium, chloride, sulfate, phosphate, bicarbonate and acetate ions on the growth rate of R. meliloti were determined. Sodium, potassium and ammonium ions had little effect on growth at concentrations of 100 mEq or less; magnesium ion inhibited growth severely at concentrations of 50 mEq (25 mM). Of the anions, chloride and sulfate appeared to have little effect while phosphate, bicarbonate, and acetate inhibited growth at concentrations of as little as 25 mEq. The addition of proline, glutamate, or betaine to cells growing in inhibitory concentrations of NaCl did not relieve the inhibition. When grown in the presence of inhibitory levels of NaCl, the intracellular concentration of glutamate but not of proline or gamma amino butyric acid increased 5-fold.     

Accumulation and localisation of sodium ions within the shoots of rice (Oryza sativa) varieties differing in salinity resistance

Oryza sativa L. (rice) is a salt-sensitive crop species which is relatively ineffective in controlling the influx of sodium and chloride ions to the shoot. Nonetheless, there is considerable varietal and individual variability in salinity resistance, much of which must derive, therefore, from differences in the fates and subsequent effects of saline ions after they have entered the plant. The destination of sodium ions within the plant has been investigated, in saline conditions, by examining the time-course of sodium ion concentrations in different leaves of four varieties and breeding lines of rice of differing salinity resistance. Radionuclide tracers were employed to study short term effects and the degree of retranslocation of these sodium ions. Sodium was not distributed uniformly but accumulated in the older leaves before the younger ones. At least some leaves were maintained at sub-lethal salt concentrations in at least the more salt resistant varieties. Radionuclide tracer studies showed that the discontinuous distribution of sodium (from leaf to leaf) is constitutive, and cannot be explained by time of exposure or differential leaf growth rates, and that significant quantities of sodium were not subsequently retranslocated, either within the plant or to the root medium.