Chloride absorption in salt-sensitive Carrizo citrange and salt-tolerant Cleopatra mandarin citrus rootstocks is linked to water use

In this work, seedlings of two citrus rootstocks, the salt-tolerant Cleopatra mandarin (Citrus reshni Hort. ex Tan.) and the salt-sensitive Carrizo citrange (Citrus sinensis [L.] Osb. x Poncirus trifoliata [L.] Raf.) were used to study the relationship between chloride and water uptake. The results indicated that net chloride uptake rates in both genotypes were alike and decreased linearly with the time of salinity exposure, although they were more rapidly reduced in the tolerant genotype. In each rootstock, chloride uptake rates paralleled the decreases in transpiration rates. When transpiration was modified, concomitant changes in leaf Cl(-) concentrations were observed. There was a high positive correlation between total chloride content per plant and total water absorbed. In addition, the data indicate that the tolerant genotype "excluded" more chloride, i.e. it absorbed lower amounts of chloride per volume of water. Cleopatra also possessed a less efficient root system for water uptake and a higher shoot-to-root ratio. The results show that, overall, chloride absorption is linked to water use and that further tolerance in Cleopatra is mostly conferred by superior root resistance to Cl(-) uptake. Therefore, it is proposed that chloride absorption and, hence, salt tolerance in citrus depends to a great extent upon water use.     

Tetraploid citrus rootstocks are more tolerant to salt stress than diploid

Citrus trees are subject to several abiotic constraints such as salinity. Providing new rootstocks more tolerant is thus a requirement. In this article, we investigated salt stress tolerance of three tetraploid rootstock genotypes when compared to their respective diploid rootstocks (Poncirus trifoliata, Carrizo citrange, Cleopatra mandarin). Plant growth, leaf fall and ion contents were investigated. At the end of the experiment, leaf fall was observed only for diploid Poncirus trifoliata plants as well as chlorosis symptoms for Poncirus trifoliata and Carrizo citrange diploid plants. The diploid Cleopatra mandarin plants growth rate was not affected by salt stress and has even been increased for tetraploid Cleopatra mandarin. Ion contents investigation has shown lower accumulations of chloride ions in leaves of the tetraploid plants when compared to diploid plants. Our results suggest that citrus tetraploid rootstocks are more tolerant to salt stress than their corresponding diploid. To cite this article: B. Saleh et al., C. R. Biologies 331 (2008).     

Morphological factors determining salt tolerance in citrus seedlings: the shoot to root ratio modulates passive root uptake of chloride ions and their accumulation in leaves

The results presented in this work were obtained with two citrus genotypes, the chloride-tolerant Cleopatra mandarin (Citrus reshni Hort. ex Tan.) and the chloride-sensitive Carrizo citrange [Citrus sinensis (L.) Osb. × Poncirus trifoliata (L.) Raf.]. The data show that chloride uptake under salinization is driven by passive forces. In both species, net rates of chloride root uptake increased linearly, without saturation, with the increase of external NaCl concentrations (30–240 mol m^–3). Uptake rates, on a μ g g root dry weight^–1 h^–1 basis, in Cleopatra and Carrizo decreased (from 38 to 21) and increased (from 21 to 35), respectively, with the increase (about three-fold) of the shoot to root ratio. With the appropriate shoot to root ratio in each genotype, it was demonstrated that at identical external doses of NaCl, Cl^– uptake rates and Cl^– xylem concentrations in the two species were very similar. Root pruning and defoliation showed that the amount of chloride taken by the plant was a function of the size of the root system, whereas leaf chloride concentration, the parameter responsible for salt damage, was dependent upon leaf biomass. Measurements of water transpiration suggested that chloride root uptake and leaf accumulation might be linked to water absorption and transpiration rates, respectively. The data indicate that plant morphology is a crucial factor determining salt-tolerance in citrus.     

Growth and root hydraulic conductivity of several citrus rootstocks under salt and polyethylene glycol stresses

The water relations responses to salt of several important citrus rootstocks such as Swingle citrumelo, sour orange, and Milam lemon have not been studied in detail before. Studies were set up to compare growth and root hydraulic properties of these rootstocks to other citrus rootstocks by exposing them to NaCl and polyethylene glycol (PEG) stresses. Seedlings of 7 citrus rootstocks were irrigated for 5 months with nutrient solutions containing NaCl or PEG that had been adjusted to osmotic potentials of -0.10, -0.20 or -0.35 MPa. The 7 rootstocks studied were sour orange (Citrus aurantium), Cleopatra mandarin (Citrus reticulata Blanco), Swingle citrumelo (C. paradisi x P. trifoliata), Carrizo citrange (C. sinensis x P. trifoliata), rough lemon (Citrus jambhiri Lush), Milam lemon (C. jambhiri hybrid), and trifoliate orange (Poncirus trifoliata [L.] Raf.). In both shoot and root growth, Cleopatra mandarin and sour orange were the least sensitive to salt, Milam and trifoliate orange were the most sensitive, and rough lemon, Swingle, and Carrizo were intermediate in sensitivity. Even though the roots were exposed to solutions of equal osmotic potentials, plant growth and root conductivity were reduced more by the PEG treatments than the corresponding NaCl treatments. At -0.10 and -0.20 MPa, shoot and root dry weights were reduced 16 to 55% by NaCl and 24 to 68% by PEG. Shoot root ratio was lowered at the higher concentrations, particularly by PEG. There was a major decrease in root conductivity caused by NaCl at -0.10 MPa (19 to 30% in sour orange and Cleopatra mandarin and 78 to 85% in trifoliate orange and Milam). Conductivity decreased more at -0.20 and -0.35 MPa, but not proportionally as much as at -0.10 MPa. Root weight per unit length increased at the higher salt levels, particularly in trifoliate orange. Water flow rate through root systems followed the same trend as root conductivity; salt affected sour orange and Cleopatra mandarin the least and trifoliate orange and Milam the most. However, reductions in fibrous root length by salt treatment differed. Root lengths of Swingle and Carrizo were least affected by salt while sour orange. Milam, and rough lemon were the most affected. Hence, even though sour orange and Cleopatra mandarin were more tolerant than the other rootstocks in terms of water flow rate or root conductivity, these 2 rootstocks showed a proportionally greater decrease in root length than Carrizo, Swingle, or trifoliate orange.     

Contrasts between Citrus species in response to salinisation: An analysis of photosynthesis and water relations for different rootstock-scion combinations

Leaf gas exchange, water relations and ion content were measured on two-year-old Valencia orange (Citrus sinensis [L.] Osbeck), Washington Navel orange (C. sinensis) and Marsh grapefruit (C. parodisi Macfad) scions budded to either Trifoliata (Poncirus infoliata [L] Raf) or Cleopatra mandarin (C. reticuLua Blanco) rootstoeks. Trees were watered with dülute nutrient solution containing either 0 or 50 _mM NaCl for 77 days. Leaf chloride concentrations (cell sap basis) were higher in all scions budded on “Trifoliata but sodium levels were lower than in equivalent foliage budded on Cleopatra mandarin rootstock. Foliar salt levels also varied according to scion. Leaves of Marsh grapefruit had higher levels of both sodium and chloride than leaves of either Valencia orange or Washington Navel orange on both rootstocks. Accumulation of sodium and chloride in salinised leaves caused a reduction in leaf osmotic potential of 0.2–1.4 MPa. and leaf water potential declined by as much as 0.5 MPa. Turgor pressure in salinised leaves was thus maintained at or above the control level. Osmotic potentials determined by psychrometry compared with pressure-volume curves were taken to imply that some accumulation of sodium or chloride in the apoplast of salinised leaves may have occurred. Despite turgor maintenance both _co2 assimilation and stomatal conductance were reduced by salinity. Following onset of leaf response to salinisation, gas exchange was impaired to a greater extent in scions budded to Cleopatra mandarin compared to those on Trifoliata. Amongst those scions. leaves of salt-treated Marsh grapefruit showed greater reductions in gas exchange than Valencia orange or Washington Navel orange budded on either rootstock. Increased sensitivity of 1Marsh grapefruit was correlated with a higher foliar sodium and chloride content in this scion. Scion differences in sensitivity of leaf gas exchange to solute concentration were independent of rootstock and appeared unrelated to leaf prolinebetaine concentrations. This implies an inherent difference between scion species with respect to salt tolerance, rather than variation in their capacity to acquire that type of compatible solute. In terms of rootstock effects, all scions proved more sensitive to salinity when budded to Cleopatra mandarin compared with Trifoliata. That response was attributed to a disproportionately higher concentration of leaf sodium in scions on Cleopatra mandarin.     

Effects of cadmium on gas exchange and phytohormone contents in citrus

The effect of increased Cd²⁺ concentrations in the watering solution on citrus physiology was studied by using two citrus genotypes, Cleopatra mandarin and Carrizo citrange. Cadmium content in roots and leaves was tested together with measurements of leaf damage, gas exchange parameters, and hormonal contents. Citrus roots efficiently retained Cd²⁺ avoiding its translocation to the shoots and Cleopatra mandarin translocated less Cd²⁺ than Carrizo. With increasing Cd²⁺ concentration all gas exchange parameters were decreased more in Carrizo than in Cleopatra mandarin. Cd-induced increases in abscisic acid and salicylic acid contents were observed in leaves but not in roots of both genotypes.     

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.     

Salinity-calcium interactions on growth and ionic concentration of Citrus plants

Two-year-old Navel orange scions (Citrus sinensis (L.) Osbeck) budded to either Cleopatra mandarin (C. reticulata) and Troyer citrange (C. sinensis × P. trifoliata) rootstocks were used in this experiment. Cleopatra manda in rootstock was considered more tolerant to salinity than Troyer citrange, and this property was attributed to a greater capacity to exclude chloride ions.Plants were grown under glasshouse conditions and supplied with nutrient solution containing either no or 45 mM NaCl. Calcium concentration was increased from 3 to 30 mM. Sodium, potassium, calcium and chloride concentrations in plant organs were analyzed after 90 days of treatment.Supplemental Ca was found to mitigate the adverse effects of salinity on plant growth, defoliation or leaf injury.Chemical analysis indicated that in plants grafted on Troyer citrange Ca restricted uptake and subsequent translocation of Na to the leaves and increased K concentration in both roots and leaves. However, in Cleopatra mandarin-grafted plants increasing Ca levels seemed to reduce transport of Na from roots to leaves, and Na accumulation in roots was associated with reduced concentration of K in this rootstock.Organ chloride analysis showed that Cl accumulation in leaves of plants grafted on both rootstocks was reduced when external Ca concentration increased, whereas Cl concentration in roots remained constant or increased. The data of distribution of Cl in plants showed that a high external Ca level increased Cl accumulation in the basal stem and roots, and reduced the transport of Cl from roots to leaves.     

Phospholipid, galactolipid and free sterol composition of fibrous roots from citrus genotypes differing in chloride exclusion ability

Abstract Fibrous roots of four citrus hybrids and parent rootstocks from which the hybrids were generated, all selected for their different Cl^? exclusion abilities, were assayed for phospholipid, galactolipid and free 4-desmethylsterol content. There was no correlation between a plant's ability to exclude Cl^? and the level of either phospholipid, galactolipid, or total free sterol in the roots of control plants. However, an inverse correlation was established between the ratio of phospholipid to free sterol in control roots and total leaf Cl^? levels of plants treated with 50 mol m^?3 NaCl for 56 d. With the exception of a significant decrease in hybrid 80-05-05, galactolipid levels were unaffected by salt treatment. Phospholipid levels were significantly increased in two parent rootstocks viz. Trifoliate orange (Poncirus trifoliata (L.). Raf.) and Carrizo citrange (Citrus sinensis (L.) Osbeck ×P. trifoliata) and one hybrid (80-02-08) but were otherwise unchanged by salt treatment. Free sterol levels were significantly increased by salt treatment in all of the better Cl^? excluders except Carrizo citrange i.e. in Rangpur lime (Citrus reticulata Blanco var. austera hybrid?), Cleopatra mandarin (Citrus reticulata Blanco) and all hybrids except 80–05–13. In all genotypes examined, salt-treatment resulted in a significant decrease in the ratio of sitosterol to stigmasterol reflecting, primarily, an increase in the stigmasterol level. The two poorer Cl^? excluders (Trifoliate orange and hybrid 80–05–13) both underwent a significant decrease in the ratio of ‘more planar’ to ‘less planar’ sterols. The inverse correlation between the phospholipid to free sterol ratio of control plants and leaf Cl^? level of salt treated plants suggests that this ratio has the potential to be used as a biochemical marker of Cl^? exclusion ability in citrus.     

Responses to flooding and drought stress by two citrus rootstock seedlings with different water-use efficiency

Leaf water relations, net gas exchange and leaf and root constituent responses to 9 days of drought stress (DS) or soil flooding were studied in 6‐month‐old seedlings of Carrizo citrange [Citrus sinensis (L.) Osb. ×Poncirus trifoliata L.; Carr] and Cleopatra mandarin (Citrus resnhi Hort. ex Tanaka; Cleo) growing in containers of native sand in the greenhouse. At the end of the drought period, both species had similar minimum stem water potentials but Cleo had higher leaf relative water content (RWC) and higher leaf osmotic potential at full turgor () than Carr. Flooding had no effect on RWC but osmotic adjustment (OA) and were higher in Cleo than in Carr. Net CO₂ assimilation rate (A_CO2) in leaves was decreased more by drought than by flooding in both species but especially in Carr. Leaf water‐use efficiency (A_CO2/transpiration) was lower in Carr and was decreased more by DS and flooding stress than in Cleo. Higher values of intercellular CO₂ concentration (C_i) in stressed plants than in control plants indicated that non‐stomatal factors including chlorophyll degradation and chlorophyll fluorescence [maximum quantum efficiency of PSII (Fv/Fm, where Fm is the maximum fluorescence and F₀, minimum fluorescence in dark‐adapted leaves)] were more important limitations on A_CO2 than stomatal conductance. In both genotypes, leaf proline was increased by drought but not by flooding, whereas both stresses increased proline in roots. Soluble sugars in leaves were increased by DS, and flooding decreased leaf sugars in Cleo. In general, DS tended to increase the concentrations of Ca, K, Mg, Na and Cl in both leaves and roots, whereas flooding tended to decrease these ions with the exception of leaf Ca in Cleo. Based on water relations and net gas exchange, Cleo was more tolerant to short‐term DS and flooding stress than Carr.     

Relationship between salt tolerance and photosynthetic machinery performance in citrus

In citrus, salt stress has been related to the build up of chloride ions in plant tissues that affect photosynthesis, growth and yield. We investigated the effects of salt stress on the stability of the photosynthetic machinery with respect to the relative salt tolerance of different citrus genotypes including: Swingle Citrumelo, Carrizo citrange, C35 citrange, Cleopatra mandarin and Forner-Alcaide #5. Under identical salt-stress conditions, Forner-Alcaide #5 and Cleopatra mandarin accumulated less chloride ions in leaves than the other genotypes and showed a better plant performance. Chlorophyll fluorescence parameters indicated severe impairments of photosynthetic activity in salt-sensitive Citrumelo and citranges but Cleopatra and Forner-Alcaide #5 were less affected. In addition, differences in photosynthetic responses between these two moderately tolerant genotypes suggested different strategies to cope with salinity. The high tolerance to salinity shown by Forner-Alcaide #5 can be associated to the ability of keeping an active photosynthetic system at elevated saline conditions whereas the tolerance of Cleopatra was linked to rapid reductions of net photosynthetic rate, stomatal conductance, performance of PSII and photosynthetic efficiency.     

Effect of NaCl concentrations in irrigation water on growth and polyamine metabolism in two citrus rootstocks with different levels of salinity tolerance

Six-months-old, uniform sized seedlings of two citrus rootstocks; Cleopatra mandarin (Citrus reshni Hort. ex Tan) and Troyer citrange (Poncirus trifoliata × Citrus sinensis) were irrigated with half-strength Hoagland nutrient solution containing 0, 40 or 80 mM NaCl for 12 weeks. Shoot height, leaf number and fresh weights of the seedlings, and relative chlorophyll contents, chlorophyll fluorescence yields (Fv/Fm), net photosynthetic and respiration rates in the leaves decreased with the increase in salinity level in the irrigation water. The decrease was greater in Troyer citrange as compared to Cleopatra mandarin. The concentrations of sugars i.e. fructose, glucose and sucrose in the leaves of Cleopatra mandarin and both leaves and roots of Troyer citrange decreased with the increase in salinity level. However, the concentrations in the roots of Cleopatra mandarin increased with the increase in salinity level. Free proline content in the leaves of Troyer citrange and root tissue of Cleopatra mandarin also increased with the increased salinity level. Among the polyamines, spermine titer increased in the leaves of both rootstocks as a response to salinity treatments. Na⁺ concentrations were higher in leaf and root tissue of Cleopatra mandarin, while that of Cl⁻ were higher in Troyer citrange.     

Physiological and biochemical adaptations of Cynodon dactylon (L.) Pers. from the Salt Range (Pakistan) to salinity stress

Naturally adapted salt tolerant populations provide a valuable material for exploring the adaptive components of salt tolerance. Under this aspect, two populations of Cynodon dactylon (L.) Pers. were subjected to salt stress in hydroponics. One was collected from a heavily salt-affected soil in the vicinity of a natural salt lake, Uchhali Lake, in the Salt Range of the Punjab province of Pakistan, and the other from a normal non-saline habitat from the Faisalabad region. The NaCl treatments in Hoagland's nutrient solution were: Control (no salt), 50, 100, 150 and 200 mM of NaCl. After 8 weeks of growth in hydroponics produced biomass, ion relations, and photosynthetic capacity were measured in the differently adapted ecotypes. In the ecotype of C. dactylon from the Salt Range, shoot dry weight was only slightly affected by varying levels of salt. However, in contrast, its root weight was markedly increased. On the other hand, the ecotype from Faisalabad (non-saline habitat) showed a marked decrease in shoot and root dry weights under saline regimes. The ecotype from the Salt Range accumulated relatively less amount of Na⁺ in the shoot than did that from Faisalabad, particularly at higher salt levels. Shoot or root K⁺ and Ca²⁺ contents varied inconsistently in both ecotypes under salt stress. All the photosynthetic parameters, leaf water potential and osmotic potential, and chlorophyll content in both ecotypes were adversely affected by salt stress, but all these physiological attributes except turgor potential and soluble sugars were less affected at high salinities in the salt tolerant ecotype from Salt Range. This ecotype accumulated significantly higher organic osmotica (total free amino acids, proline, total soluble proteins, and total soluble sugars) under saline conditions than its intolerant counterpart. Overall, the salt tolerant ecotype of C. dactylon from the Salt Range showed high salt tolerance due to its restricted uptake of Na⁺ accompanied by an increased uptake of K⁺ and Ca²⁺ in the roots as well as shoot due to its higher photosynthetic capacity and accumulation of organic osmotica such as free amino acids and proline under saline conditions.     

Mechanism of paraquat tolerance in perennial ryegrass

Abstract The mechanism of paraquat tolerance was investigated in lines of perennial ryegrass (Lolium perenne L.) which had been selected for resistance to the herbicide. Uptake, metabolism and translocation of paraquat were studied. Susceptible cultivars and a tolerant line were not found to differ in uptake of radioactive paraquat applied to the leaf surface or supplied to the cut ends of excised leaves. Distribution of herbicide within leaf tissue was similar in tolerant and susceptible plants and no metabolites of ¹⁴C-paraquat were detected in tolerant or susceptible plants treated with sub-lethal concentrations of the herbicide. Autoradiography and quantitative determinations showed much variation in translocation of ¹⁴C-paraquat out of treated leaves of intact plants, but the variation was not related to the degree of susceptibility to the herbicide. It is concluded that paraquat tolerance in perennial ryegrass is unlikely to depend upon reduced uptake, enhanced metabolism or altered translocation of the herbicide.     

Alleviation of salt stress in citrus seedlings inoculated with arbuscular mycorrhizal fungi depends on the rootstock salt tolerance

Seedlings of Cleopatra mandarin (Citrus reshni Hort. ex Tan.) and Alemow (Citrus macrophylla Wester) were inoculated with a mixture of AM fungi (Rhizophagus irregularis and Funneliformis mosseae) (+AM), or left non-inoculated (−AM). From forty-five days after fungal inoculation onwards, half of +AM or −AM plants were irrigated with nutrient solution containing 50 mM NaCl. Three months later, AM significantly increased plant growth in both Cleopatra mandarin and Alemow rootstocks. Plant growth was higher in salinized +AM plants than in non-salinized −AM plants, demonstrating that AM compensates the growth limitations imposed by salinity. Whereas AM-inoculated Cleopatra mandarin seedlings had a very good response under saline treatment, inoculation in Alemow did not alleviate the negative effect of salinity. The beneficial effect of mycorrhization is unrelated with protection against the uptake of Na or Cl and the effect of AM on these ions did not explain the different response of rootstocks. This response was related with the nutritional status since our findings confirm that AM fungi can alter host responses to salinity stress, improving more the P, K, Fe and Cu plant nutrition in Cleopatra mandarin than in Alemow plants. AM inoculation under saline treatments also increased root Mg concentration but it was higher in Cleopatra mandarin than in Alemow. This could explain why AM fungus did not completely recovered chlorophyll concentrations in Alemow and consequently it had lower photosynthesis rate than control plants. AM fungi play an essential role in citrus rootstock growth and biomass production although the intensity of this response depends on the rootstock salinity tolerance.     

Salinity tolerance of 'Valencia' orange trees on rootstocks with contrasting salt tolerance is not improved by moderate shade

The effects of shading in combination with salinity treatments were studied in citrus trees on two rootstocks with contrasting salt tolerance to determine if shading could reduce the negative effects of salinity stress. Well-nourished 2-year-old 'Valencia' orange trees grafted on Cleopatra mandarin (Cleo, relatively salt tolerant) or Carrizo citrange (Carr, relatively salt sensitive), were grown either under a 50% shade cloth or left unshaded in full sunlight. Half the trees received no salinity treatment and half were salinized with 50 mM Cl- during two 9 week salinity periods in the spring and autumn interrupted by an 11 week rainy period. The shade treatment reduced midday leaf temperature and leaf-to-air vapour pressure deficit regardless of salinity treatments. In non-salinized trees, shade increased midday CO2 assimilation rate (A(CO2)) and stomatal conductance, but had no effect on leaf transpiration (E(lf)). Shade also increased leaf chlorophyll and photosynthetic water use efficiency (A(CO2)/E(lf)) in leaves on both rootstocks and increased total plant dry weight in Cleo. The salinity treatment reduced leaf growth and leaf gas exchange parameters. Shade decreased Cl- concentrations in leaves of salinized Carr trees, but had no effect on leaf or root Cl- of trees on Cleo. There were no significant differences in leaf gas exchange parameters of shaded and unshaded salinized plants but the growth reduction from salinity stress was actually greater for shaded than for unshaded trees. Shaded trees on both rootstocks had higher leaf Na+ than unshaded trees after the first salinity period, and this shade-induced elevated leaf Na+ persisted after the second salinity period in trees on Carr. Thus, shading did not alleviate the negative effects of salinity on growth and Na+ accumulation.     

Anatomical and physiological characteristics relating to ionic relations in some salt tolerant grasses from the Salt Range,Pakistan

Populations of three salt tolerant forage grasses (Cynodon dactylon, Imperata cylindrica, and Sporobolus arabicus) were collected from the salt-affected soils of the Salt Range and normal non-saline soils of the Faisalabad region to assess their mechanism of adaptation to saline stress by determining ion relations and some specific anatomical modifications. The population of S. arabicus from the Salt Range showed increased growth (root and shoot length, and root and shoot dry weights) under saline conditions. Salt tolerance in this species was related to structural modifications such as increased area of root, stem, leaf blade, and leaf sheath for toxic ion accumulation, increased vesicular hair density in leaves and aerenchyma formation in leaf sheath for ion exclusion. Uptake of toxic ions was high in the Salt Range population of C. dactylon and salt tolerance was related to ion exclusion through specific leaf structural modifications such as vesicular hairs. Salt tolerance in the Salt Range population of I. cylindrica was mainly associated with restricted uptake of toxic Na⁺ and Cl⁻ at root level, and accumulation of toxic ions via increased succulence in leaf blades and leaf sheaths in addition to some excretion of toxic ions through leaf sheath aerenchyma.     

Variation in growth and ion accumulation between two selected populations of Trifolium repens L. differing in salt tolerance

Two divergent populations of T. repens cv. Haifa developed from two generations of recurrent selection for shoot chloride concentration, were grown in the greenhouse at 0 and 40 mol m^–3 NaCl. Over two harvest cycles at 40 mol m^–3 NaCl, the population selected for a low concentration of chloride in the shoot maintained a significantly lower chloride and sodium concentration compared with those plants selected for a high shoot chloride concentration. The distribution of chloride in the shoots was further examined in a subsample of plants from both populations. In all plants, concentrations of chloride were lower in the expanding and fully expanded leaves than in the older leaf tissue or petioles.While there were no significant differences in the photosynthetic rates between lines, shoot yields and relative leaf expansion rates were higher in the low chloride population. Plant death was greater in plants selected for high shoot chloride. These results suggest that selections based on measurements of low shoot chloride concentrations may be successful in developing a cultivar of T. repens with improved salt tolerance.