Change in uptake, transport and accumulation of ions in Nerium oleander (rosebay) as affected by different nitrogen sources and salinity

Background and Aims

The source of nitrogen plays an important role in salt tolerance of plants. In this study, the effects of NaCl on net uptake, accumulation and transport of ions were investigated in Nerium oleander with ammonium or nitrate as the nitrogen source in order to analyse differences in uptake and cycling of ions within plants.

Methods

Plants were grown in a greenhouse in hydroponics under different salt treatments (control vs. 100 mm NaCl) with ammonium or nitrate as the nitrogen source, and changes in ion concentration in plants, xylem sap exuded from roots and stems, and phloem sap were determined.

Key Results

Plant weight, leaf area and photosynthetic rate showed a higher salt tolerance of nitrate-fed plants compared with that of ammonium-fed plants. The total amount of Na⁺ transported in the xylem in roots, accumulated in the shoot and retranslocated in the phloem of ammonium-fed plants under salt treatment was 1·8, 1·9 and 2·7 times more, respectively, than that of nitrate-treated plants. However, the amount of Na⁺ accumulated in roots in nitrate-fed plants was about 1·5 times higher than that in ammonium-fed plants. Similarly, Cl⁻ transport via the xylem to the shoot and its retranslocation via the phloem (Cl⁻ cycling) were far greater with ammonium treatment than with nitrate treatment under conditions of salinity. The uptake and accumulation of K⁺ in shoots decreased more due to salinity in ammonium-fed plants compared with nitrate-fed plants. In contrast, K⁺ cycling in shoots increased due to salinity, with higher rates in the ammonium-treated plants.

Conclusions

The faster growth of nitrate-fed plants under conditions of salinity was associated with a lower transport and accumulation of Na⁺ and Cl⁻ in the shoot, whereas in ammonium-fed plants accumulation and cycling of Na⁺ and Cl⁻ in shoots probably caused harmful effects and reduced growth of plants.Key words: Mineral cycling, Nerium oleander, nitrogen source, salinity, xylem and phloem transport     

Tolerance of combined submergence and salinity in the halophytic stem-succulent Tecticornia pergranulata

Background and Aims

Habitats occupied by many halophytes are not only saline, but are also prone to flooding. Few studies have evaluated submergence tolerance in halophytes.

Methods

Responses to submergence, at a range of salinity levels, were studied for the halophytic stem-succulent Tecticornia pergranulata subsp. pergranulata (syn. Halosarcia pergranulata subsp. pergranulata). Growth and total sugars in succulent stems were assessed as a function of time after submergence. Underwater net photosynthesis, dark respiration, total sugars, glycinebetaine, Na⁺, Cl⁻ and K⁺, in succulent stems, were assessed in a NaCl dose-response experiment.

Key Results

Submerged plants ceased to grow, and tissue sugars declined. Photosynthesis by succulent stems was reduced markedly when underwater, as compared with in air. Capacity for underwater net photosynthesis (P_N) was not affected by 10–400 mm NaCl, but it was reduced by 30 % at 800 mm. Dark respiration, underwater, increased in succulent stems at 200–800 mm NaCl, as compared with those at 10 mm NaCl. On an ethanol-insoluble dry mass basis, K⁺ concentration in succulent stems of submerged plants was equal to that in drained controls, across all NaCl treatments. Na⁺ and Cl⁻ concentrations, however, were elevated in stems of submerged plants, but so was glycinebetaine. Submerged stems increased in succulence, so solutes would have been ‘diluted’ on a tissue-water basis.

Conclusions

Tecticornia pergranulata tolerates complete submergence, even in waters of high salinity. A ‘quiescence response’, i.e. no shoot growth, would conserve carbohydrates, but tissue sugars still declined with time. A low K⁺ : Na⁺ ratio, typical for tissues of succulent halophytes, was tolerated even during prolonged submergence, as evidenced by maintenance of underwater P_N at up to 400 mm NaCl. Underwater P_N provides O₂ and sugars, and thus should enhance survival of submerged plants.Key words: Flooding, halophyte, Halosarcia pergranulata, inundation, inland salt marsh, respiration, Salicornioideae, salt lake, submergence–salinity interaction, tissue solutes, underwater net photosynthesis     

Ecophysiological response of Crambe maritima to airborne and soil-borne salinity

Background and Aims

There is a need to evaluate the salt tolerance of plant species that can be cultivated as crops under saline conditions. Crambe maritima is a coastal plant, usually occurring on the driftline, with potential use as a vegetable crop. The aim of this experiment was to determine the growth response of Crambe maritima to various levels of airborne and soil-borne salinity and the ecophysiological mechanisms underlying these responses.

Methods

In the greenhouse, plants were exposed to salt spray (400 mm NaCl) as well as to various levels of root-zone salinity (RZS) of 0, 50, 100, 200 and 300 mm NaCl during 40 d. The salt tolerance of Crambe maritima was assessed by the relative growth rate (RGR) and its components. To study possible salinity effects on the tissue and cellular level, the leaf succulence, tissue Na⁺ concentrations, Na⁺ : K⁺ ratio, net K⁺/Na⁺ selectivity, N, P, K⁺, Ca²⁺, Mg²⁺, proline, soluble sugar concentrations, osmotic potential, total phenolics and antioxidant capacity were measured.

Key Results

Salt spray did not affect the RGR of Crambe maritima. However, leaf thickness and leaf succulence increased with salt spray. Root zone salinities up to 100 mm NaCl did not affect growth. However, at 200 mm NaCl RZS the RGR was reduced by 41 % compared with the control and by 56 % at 300 mm NaCl RZS. The reduced RGR with increasing RZS was largely due to the reduced specific leaf area, which was caused by increased leaf succulence as well as by increased leaf dry matter content. No changes in unit leaf rate were observed but increased RZS resulted in increased Na⁺ and proline concentrations, reduced K⁺, Ca²⁺ and Mg²⁺ concentrations, lower osmotic potential and increased antioxidant capacity. Proline concentrations of the leaves correlated strongly (r = 0·95) with RZS concentrations and not with plant growth.

Conclusions

Based on its growth response, Crambe maritima can be classified as a salt spray tolerant plant that is sensitive to root zone salinities exceeding 100 mm NaCl.     

A novel protein kinase involved in Na+ exclusion revealed from positional cloning

Salinity is a major abiotic stress which affects crop plants around the world, resulting in substantial loss of yield and millions of dollars of lost revenue. High levels of Na⁺ in shoot tissue have many adverse effects and, crucially, yield in cereals is commonly inversely proportional to the extent of shoot Na⁺ accumulation. We therefore need to identify genes, resistant plant cultivars and cellular processes that are involved in salinity tolerance, with the goal of introducing these factors into commercially available crops. Through the use of an Arabidopsis thaliana mapping population, we have identified a highly significant quantitative trait locus (QTL) linked to Na⁺ exclusion. Fine mapping of this QTL identified a protein kinase (AtCIPK16), related to AtSOS2, that was significantly up‐regulated under salt stress. Greater Na⁺ exclusion was associated with significantly higher root expression of AtCIPK16, which is due to differences in the gene's promoter. Constitutive overexpression of the gene in Arabidopsis leads to plants with significant reduction in shoot Na⁺ and greater salinity tolerance. amiRNA knock‐downs of AtCIPK16 in Arabidopsis show a negative correlation between the expression levels of the gene and the amount of shoot Na⁺. Transgenic barley lines overexpressing AtCIPK16 show increased salinity tolerance.     

Improving crop salt tolerance using transgenic approaches: An update and physiological analysis

Salinization of land is likely to increase due to climate change with impact on agricultural production. Since most species used as crops are sensitive to salinity, improvement of salt tolerance is needed to maintain global food production. This review summarises successes and failures of transgenic approaches in improving salt tolerance in crop species. A conceptual model of coordinated physiological mechanisms in roots and shoots required for salt tolerance is presented. Transgenic plants overexpressing genes of key proteins contributing to Na⁺ ‘exclusion’ (PM-ATPases with SOS1 antiporter, and HKT1 transporter) and Na⁺ compartmentation in vacuoles (V-H⁺ATPase and V-H⁺PPase with NHX antiporter), as well as two proteins potentially involved in alleviating water deficit during salt stress (aquaporins and dehydrins), were evaluated. Of the 51 transformations, with gene(s) involved in Na⁺ ‘exclusion’ or Na⁺ vacuolar compartmentation that contained quantitative data on growth and include a non-saline control, 48 showed improvements in salt tolerance (less impact on plant mass) of transgenic plants, but with only two tested in field conditions. Of these 51 transformations, 26 involved crop species. Tissue ion concentrations were altered, but not always in the same way. Although glasshouse data are promising, field studies are required to assess crop salinity tolerance.     

Tolerance of Hordeum marinum accessions to O2 deficiency, salinity and these stresses combined

Background and Aims

When root-zone O₂ deficiency occurs together with salinity, regulation of shoot ion concentrations is compromised even more than under salinity alone. Tolerance was evaluated amongst 34 accessions of Hordeum marinum, a wild species in the Triticeae, to combined salinity and root-zone O₂ deficiency. Interest in H. marinum arises from the potential to use it as a donor for abiotic stress tolerance into wheat.

Methods

Two batches of 17 H. marinum accessions, from (1) the Nordic Gene Bank and (2) the wheat belt of Western Australia, were exposed to 0·2 or 200 mol m⁻³ NaCl in aerated or stagnant nutrient solution for 28–29 d. Wheat (Triticum aestivum) was included as a sensitive check species. Growth, root porosity, root radial O₂ loss (ROL) and leaf ion (Na⁺, K⁺, Cl⁻) concentrations were determined.

Key Results

Owing to space constraints, this report is focused mainly on the accessions from the Nordic Gene Bank. The 17 accessions varied in tolerance; relative growth rate was reduced by 2–38 % in stagnant solution, by 8–42 % in saline solution (aerated) and by 39–71 % in stagnant plus saline treatment. When in stagnant solution, porosity of adventitious roots was 24–33 %; salinity decreased the root porosity in some accessions, but had no effect in others. Roots grown in stagnant solution formed a barrier to ROL, but variation existed amongst accessions in apparent barrier ‘strength’. Leaf Na⁺ concentration was 142–692 µmol g⁻¹ d. wt for plants in saline solution (aerated), and only increased to 247–748 µmol g⁻¹ d. wt in the stagnant plus saline treatment. Leaf Cl⁻ also showed only small effects of stagnant plus saline treatment, compared with saline alone. In comparison with H. marinum, wheat was more adversely affected by each stress alone, and particularly when combined; growth reductions were greater, adventitious root porosity was 21 %, it lacked a barrier to ROL, leaf K⁺ declined to lower levels, and leaf Na⁺ and Cl⁻ concentrations were 3·1–9-fold and 2·8–6-fold higher, respectively, in wheat.

Conclusions

Stagnant treatment plus salinity reduced growth more than salinity alone, or stagnant alone, but some accessions of H. marinum were still relatively tolerant of these combined stresses, maintaining Na⁺ and Cl⁻ ‘exclusion’ even in an O₂-deficient, saline rooting medium.Key words: Aerenchyma, combined salinity and waterlogging, leaf Cl⁻, leaf K⁺, leaf Na⁺, radial O₂ loss, salt tolerance, salinity–waterlogging interaction, sea barleygrass, waterlogging tolerance, wheat, wild Triticeae     

A teosinte-derived allele of an HKT1 family sodium transporter improves salt tolerance in maize

The sodium cation (Na⁺) is the predominant cation with deleterious effects on crops in salt-affected agricultural areas. Salt tolerance of crop can be improved by increasing shoot Na⁺ exclusion. Therefore, it is crucial to identify and use genetic variants of various crops that promote shoot Na⁺ exclusion. Here, we show that a HKT1 family gene ZmNC3 (Zea mays L. Na⁺ Content 3; designated ZmHKT1;2) confers natural variability in shoot-Na⁺ accumulation and salt tolerance in maize. ZmHKT1;2 encodes a Na⁺-preferential transporter localized in the plasma membrane, which mediates shoot Na⁺ exclusion, likely by withdrawing Na⁺ from the root xylem flow. A naturally occurring nonsynonymous SNP (SNP947-G) increases the Na⁺ transport activity of ZmHKT1;2, promoting shoot Na⁺ exclusion and salt tolerance in maize. SNP947-G first occurred in the wild grass teosinte (at a allele frequency of 43%) and has become a minor allele in the maize population (allele frequency 6.1%), suggesting that SNP947-G is derived from teosinte and that the genomic region flanking SNP947 likely has undergone selection during domestication or post-domestication dispersal of maize. Moreover, we demonstrate that introgression of the SNP947-G ZmHKT1;2 allele into elite maize germplasms reduces shoot Na⁺ content by up to 80% and promotes salt tolerance. Taken together, ZmNC3/ZmHKT1;2 was identified as an important QTL promoting shoot Na⁺ exclusion, and its favourable allele provides an effective tool for developing salt-tolerant maize varieties.     

Control of xylem Na+ loading and transport to the shoot in rice and barley as a determinant of differential salinity stress tolerance

Control of xylem Na⁺ loading has often been named as the essential component of salinity tolerance mechanism. However, it is less clear to what extent the difference in this trait may determine differential salinity tolerance between species. In this study, barley (Hordeum vulgare L. cv. CM72) and rice (Oryza sativa L. cv. Dongjin) plants were grown under two levels of salinity. Na⁺ and K⁺ concentrations in the xylem sap, and shoot and root tissues were measured at different time points after stress onset. Salt‐exposed rice plants prevented xylem Na⁺ loading for several days, but failed to control this process in the longer term, ultimately resulting in a massive Na⁺ shoot loading. Barley plants quickly increased xylem Na⁺ concentration and its delivery to the shoot (most likely for the purpose of osmotic adjustment) but were able to reduce this process later on, keeping most of accumulated Na⁺ in the root, thus maintaining non‐toxic shoot Na⁺ level. Rice plants increased shoot K⁺ concentration, while barley plants maintained higher root K⁺ concentration. Control of xylem Na⁺ loading is remarkably different between rice and barley; this difference may differentiate the extent of the salinity tolerance between species. This trait should be investigated in more detail to be used in the breeding programs aimed to improve salinity tolerance in crops.     

Saline-induced changes of epicuticular waxy layer on the Puccinellia tenuiflora and Oryza sativa leave surfaces

Background

The epicuticular waxy layer of plant leaves enhances the extreme environmental stress tolerance. However, the relationship between waxy layer and saline tolerance was not established well. The epicuticular waxy layer of rice (Oryza sativa L.) was studied under the NaHCO₃ stresses. In addition, strong saline tolerance Puccinellia tenuiflora was chosen for comparative studies.

Results

Scanning electron microscope (SEM) images showed that there were significant changes in waxy morphologies of the rice epicuticular surfaces, while no remarkable changes in those of P. tenuiflora epicuticular surfaces. The NaHCO₃-induced morphological changes of the rice epicuticular surfaces appeared as enlarged silica cells, swollen corns-shapes and leaked salt columns under high stress. Energy dispersive X-ray (EDX) spectroscopic profiles supported that the changes were caused by significant increment and localization of [Na⁺] and [Cl⁻] in the shoot. Atomic absorption spectra showed that [Na⁺]_shoot/[Na⁺]_root for P. tenuiflora maintained stable as the saline stress increased, but that for rice increased significantly.

Conclusion

In rice, NaHCO₃ stress induced localization and accumulation of [Na⁺] and [Cl⁻] appeared as the enlarged silica cells (MSC), the swollen corns (S-C), and the leaked columns (C), while no significant changes in P. tenuiflora.     

Response to non-uniform salinity in the root zone of the halophyte Atriplex nummularia: growth, photosynthesis, water relations and tissue ion concentrations

Background and Aims

Soil salinity is often heterogeneous, yet the physiology of halophytes has typically been studied with uniform salinity treatments. An evaluation was made of the growth, net photosynthesis, water use, water relations and tissue ions in the halophytic shrub Atriplex nummularia in response to non-uniform NaCl concentrations in a split-root system.

Methods

Atriplex nummularia was grown in a split-root system for 21 d, with either the same or two different NaCl concentrations (ranging from 10 to 670 mm), in aerated nutrient solution bathing each root half.

Key Results

Non-uniform salinity, with high NaCl in one root half (up to 670 mm) and 10 mm in the other half, had no effect on shoot ethanol-insoluble dry mass, net photosynthesis or shoot pre-dawn water potential. In contrast, a modest effect occurred for leaf osmotic potential (up to 30 % more solutes compared with uniform 10 mm NaCl treatment). With non-uniform NaCl concentrations (10/670 mm), 90 % of water was absorbed from the low salinity side, and the reduction in water use from the high salinity side caused whole-plant water use to decrease by about 30 %; there was no compensatory water uptake from the low salinity side. Leaf Na⁺ and Cl⁻ concentrations were 1·9- to 2·3-fold higher in the uniform 670 mm treatment than in the 10/670 mm treatment, whereas leaf K⁺ concentrations were 1·2- to 2·0-fold higher in the non-uniform treatment.

Conclusions

Atriplex nummularia with one root half in 10 mm NaCl maintained net photosynthesis, shoot growth and shoot water potential even when the other root half was exposed to 670 mm NaCl, a concentration that inhibits growth by 65 % when uniform in the root zone. Given the likelihood of non-uniform salinity in many field situations, this situation would presumably benefit halophyte growth and physiology in saline environments.Key words: Split-root system, salinity heterogeneity, root zone heterogeneity, water potential, water use, stomatal conductance, saltbush, leaf ions, photosynthesis, NaCl     

How much does agriculture depend on pollinators? Lessons from long-term trends in crop production

Background and Aims

Productivity of many crops benefits from the presence of pollinating insects, so a decline in pollinator abundance should compromise global agricultural production. Motivated by the lack of accurate estimates of the size of this threat, we quantified the effect of total loss of pollinators on global agricultural production and crop production diversity. The change in pollinator dependency over 46 years was also evaluated, considering the developed and developing world separately.

Methods

Using the extensive FAO dataset, yearly data were compiled for 1961–2006 on production and cultivated area of 87 important crops, which we classified into five categories of pollinator dependency. Based on measures of the aggregate effect of differential pollinator dependence, the consequences of a complete loss of pollinators in terms of reductions in total agricultural production and diversity were calculated. An estimate was also made of the increase in total cultivated area that would be required to compensate for the decrease in production of every single crop in the absence of pollinators.

Key Results

The expected direct reduction in total agricultural production in the absence of animal pollination ranged from 3 to 8 %, with smaller impacts on agricultural production diversity. The percentage increase in cultivated area needed to compensate for these deficits was several times higher, particularly in the developing world, which comprises two-thirds of the land devoted to crop cultivation globally. Crops with lower yield growth tended to have undergone greater expansion in cultivated area. Agriculture has become more pollinator-dependent over time, and this trend is more pronounced in the developing than developed world.

Conclusions

We propose that pollination shortage will intensify demand for agricultural land, a trend that will be more pronounced in the developing world. This increasing pressure on supply of agricultural land could significantly contribute to global environmental change.Key words: Agricultural production, biotic pollination, crop diversity, cultivated area, developed world, developing world, FAO, randomization     

Plant salt tolerance: adaptations in halophytes

Background Most of the water on Earth is seawater, each kilogram of which contains about 35 g of salts, and yet most plants cannot grow in this solution; less than 0·2 % of species can develop and reproduce with repeated exposure to seawater. These ‘extremophiles’ are called halophytes.Scope Improved knowledge of halophytes is of importance to understanding our natural world and to enable the use of some of these fascinating plants in land re-vegetation, as forages for livestock, and to develop salt-tolerant crops. In this Preface to a Special Issue on halophytes and saline adaptations, the evolution of salt tolerance in halophytes, their life-history traits and progress in understanding the molecular, biochemical and physiological mechanisms contributing to salt tolerance are summarized. In particular, cellular processes that underpin the ability of halophytes to tolerate high tissue concentrations of Na⁺ and Cl⁻, including regulation of membrane transport, their ability to synthesize compatible solutes and to deal with reactive oxygen species, are highlighted. Interacting stress factors in addition to salinity, such as heavy metals and flooding, are also topics gaining increased attention in the search to understand the biology of halophytes.Conclusions Halophytes will play increasingly important roles as models for understanding plant salt tolerance, as genetic resources contributing towards the goal of improvement of salt tolerance in some crops, for re-vegetation of saline lands, and as ‘niche crops’ in their own right for landscapes with saline soils.     

Osmoregulation in Plants: Implications for Agriculture

Drought and salinity stress are the major causes of historicand modern agricultural productivity losses throughout the world.Both drought and salinity result in osmotic stress that maylead to inhibition of growth. Salinity causes additional iontoxicity effects mainly through perturbations in protein andmembrane structure. In contrast to animals, which rely on Na⁺/K⁺-ATPasesfor the expulsion of osmotica, plants rely on plasma membraneand endosomal ATPase activities to generate proton gradientsto drive ion extrusion and intracellular sequestration. Consequently,most angiosperms, including all major crop species, have a diminishedcapacity for Na⁺ transport and tolerance to high salinity. Newinsights into the molecular mechanisms of Na⁺/K⁺ discrimination,Na⁺ extrusion and compartmentation, water transport, and osmolytebiosynthesis and function have led to genetically engineeredplants with improved salt, drought, and cold tolerance. A deeperunderstanding of the complex signal transduction and regulatoryresponses to osmotic stress promises novel strategies for improvingsalinity and drought tolerance that will be of practical benefitto agriculture.     

Salt sensitivity in chickpea is determined by sodium toxicity

Main conclusion

Salt sensitivity in chickpea is determined by Na⁺ toxicity, whereas relatively high leaf tissue concentrations of Cl^? were tolerated, and the osmotic component of 60-mM NaCl was not detrimental.Chickpea (Cicer arietinum L.) is sensitive to salinity. This study dissected the responses of chickpea to osmotic and ionic components (Na⁺ and/or Cl^?) of salt stress. Two genotypes with contrasting salt tolerances were exposed to osmotic treatments (?0.16 and ?0.29 MPa), Na⁺-salts, Cl^?-salts, or NaCl at 0, 30, or 60 mM for 42 days and growth, tissue ion concentrations and leaf gas-exchange were assessed. The osmotic treatments and Cl^?-salts did not affect growth, whereas Na⁺-salts and NaCl treatments equally impaired growth in either genotype. Shoot Na⁺ and Cl^? concentrations had markedly increased, whereas shoot K⁺ had declined in the NaCl treatments, but both genotypes had similar shoot concentrations of each of these individual ions after 14 and 28 days of treatments. Genesis836 achieved higher net photosynthetic rate (64–84 % of control) compared with Rupali (35–56 % of control) at equivalent leaf Na⁺ concentrations. We conclude that (1) salt sensitivity in chickpea is determined by Na⁺ toxicity, and (2) the two contrasting genotypes appear to differ in ‘tissue tolerance’ of high Na⁺. This study provides a basis for focus on Na⁺ tolerance traits for future varietal improvement programs for salinity tolerance in chickpea.

     

Ecosystem changes in Galápagos highlands by the invasive tree Cinchona pubescens

Background and aims

Salinity is an increasing problem for agricultural production worldwide. Understanding how Na⁺ enters plants is important if reducing Na⁺ influx, a key component of the regulation of Na⁺ accumulation in plants and improving salt tolerance of crop plants, is to be achieved. Our previous work indicated that two distinct low-affinity Na⁺ uptake pathways exist in the halophyte Suaeda maritima. Here, we report the external NaCl concentration at which uptake switches from pathway 1 to pathway 2 and the kinetics of the interaction between external K⁺ concentration and Na⁺ uptake and accumulation in S. maritima in order to determine the roles of K⁺ transporters or channels in low-affinity Na⁺ uptake.

Methods

Na⁺ influx, Na⁺ and K⁺ accumulations in S. maritima exposed to various concentrations of NaCl (0–200 mM) were analyzed in the absence and presence of the inhibitors TEA and Ba⁺ (5 mM TEA or 3 mM Ba²⁺) or KCl (0, 10 or 50 mM).

Results

Our earlier proposal was confirmed and extended that there are two distinct low-affinity Na⁺ uptake pathways in S. maritima: pathway 1 might be mediated by a HKT-type transporter under low salinity conditions and pathway 2 by an AKT1-type channel or a KUP/HAK/KT type transporter under high salinity conditions. The external NaCl concentration at which two distinct low-affinity Na⁺ uptake switches from pathway 1 to pathway 2, the ‘turning point’, is between 90 and 95 mM. Over a short period (12 h) of Na⁺ and K⁺ treatments, a low concentration of K⁺ (10 mM) facilitated Na⁺ uptake by S. maritima under high salinity (100–200 mM NaCl), whether or not the plants had been subjected to a longer (3 d) period of K⁺ starvation. The kinetics suggests that low concentration of K⁺ (10 mM) might activate AKT1-type channels or KUP/HAK/KT-type transporters under high salinity (100–200 mM NaCl).

Conclusions

The turning-point of external NaCl concentrations for the two low-affinity Na⁺ uptake pathways in Suaeda maritima is between 90 and 95 mM. A low concentration of K⁺ (10 mM) might activate AKT1 or KUP/HAK/KT and facilitate Na⁺ uptake under high salinity (100–200 mM NaCl). The kinetics of K⁺ on Na⁺ uptake and accumulation in S maritima are also consistent with there being two low-affinity Na⁺ uptake pathways.