盐胁迫对二穗短柄草幼苗生长及不同器官中盐离子稳态的影响

采用4种浓度的NaCl溶液(50、100、150、200 mmol/L)处理二穗短柄草和拟南芥(对照)幼苗,测定不同浓度盐胁迫下2种植物幼苗的生长指标和离子分布,以探讨二穗短柄草在盐胁迫下主要阳离子平衡机制.结果表明:(1)盐胁迫显著抑制二穗短柄草根叶的生物量积累.(2)根冠比数据显示,在盐胁迫条件下二穗短柄草能够更好地维系地下部分的生物量积累.(3)在4种浓度盐胁迫下,二穗短柄草叶中Na+含量低于根系,而且K+、Cl-含量和K+/Na+比值始终高于根系,说明在二穗短柄草中Na+从地下到地上的转运受到抑制,但对Cl-的转运缺乏有效的调控.(4)回归分析发现,盐胁迫下二穗短柄草和拟南芥根部Na+与K+含量变化呈正相关关系,而在叶部则不相关,说明二穗短柄草和拟南芥Na+与K+在根部具有相同的离子通道,而在叶部却具有各自独立的转运途径.     

高盐生境中乌拉尔甘草盐离子分泌与积存格局的研究

采用植物水培方法,以乌拉尔甘草为研究材料,用不同浓度(0、80、160、320mmol·L~(-1))NaCl溶液胁迫处理乌拉尔甘草幼苗3周后,分析其叶片表面盐离子(K~+、Ca~(2+)、Na+)分泌速率的差异,并采集盐化低地草甸重盐土生境中2年生乌拉尔甘草植株,应用ICP-AES测定其不同部位(根、根状茎、茎、老叶和幼叶)中的盐离子(K~+、Na~+、Ga~(2+)、Mg~(2+))含量,探究盐离子在乌拉尔甘草叶片上的分泌格局以及盐离子在植株体内的积存格局,为完善甘草耐盐机理的研究提供依据。结果显示:(1)随着盐胁迫浓度的升高,乌拉尔甘草叶片上K~+、Ca~(2+)、Na+的分泌速率均呈增加趋势,且Na~+的分泌速率远远大于Ca~(2+)和K+的分泌速率。(2)在乌拉尔甘草各部位中,K+的积存量从大到小依次为:幼叶根根状茎茎老叶;Na~+在各个部位的积存量都十分有限,且无论地下部分还是地上部分,差异均不大;Ca~(2+)积存量由大到小依次为:老叶幼叶茎根状茎根,且老叶中Ca~(2+)的积存量显著高于其它部位。研究认为,在重盐碱地生境中,K+主要积存在幼叶中,Ga~(2+)主要积存在老叶中,植株体内各个部位Na~+的积存量很低,乌拉尔甘草表现出明显的拒Na现象;叶片分泌的主要盐离子为Na~+;乌拉尔甘草通过泌盐的方式将Na+排出体外,从而有效降低Na~+在体内的积存,这是其能够在重盐碱地生存生长的重要原因。     

Root growth inhibition by NH4+ in Arabidopsis is mediated by the root tip and is linked to NH4+ efflux and GMPase activity

Root growth in higher plants is sensitive to excess ammonium (NH₄⁺). Our study shows that contact of NH₄⁺ with the primary root tip is both necessary and sufficient to the development of arrested root growth under NH₄⁺ nutrition in Arabidopsis. We show that cell elongation and not cell division is the principal target in the NH₄⁺ inhibition of primary root growth. Mutant and expression analyses using DR5:GUS revealed that the growth inhibition is furthermore independent of auxin and ethylene signalling. NH₄⁺ fluxes along the primary root, measured using the Scanning Ion‐selective Electrode Technique, revealed a significant stimulation of NH₄⁺ efflux at the elongation zone following treatment with elevated NH₄⁺, coincident with the inhibition of root elongation. Stimulation of NH₄⁺ efflux and inhibition of cell expansion were significantly more pronounced in the NH₄⁺‐hypersensitive mutant vtc1‐1, deficient in the enzyme GDP‐mannose pyrophosphorylase (GMPase). We conclude that both restricted transmembrane NH₄⁺ fluxes and proper functioning of GMPase in roots are critical to minimizing the severity of the NH₄⁺ toxicity response in Arabidopsis.     

Salt-Induced Hypodermal Transfer Cells in Roots of Prosopis farcta and Ion Distribution within Young Plants

Prosopis farcta was grown on hydroculture with additions of 0.5, 10, 50, and 100 mM NaCl and without salt treatment. In plants from a 0.5 mM NaCl treatment, Cl^? was taken up into stems and leaves, but Na⁺ was withheld from the shoot. At 10 mM NaCl, shoot K⁺ concentration was below that of the control; Na⁺ and Cl^? were taken up to stems and cotyledons in nearly equimolar amounts. However, in the leaves, Na⁺ concentrations were only half of those of Cl^?. With increasing salt stress, Na⁺ and Cl^? were transported to the shoot, but kept at relatively low levels in the roots. Na⁺/ K⁺ ratios in roots did not increase proportionally to those in the solution. At an external Na⁺/K⁺ of > 5 and a root Na⁺/K⁺ of >1 (10 mM NaCl treatment), K⁺ selectivity was induced which rose exponentially with increasing salt stress; and cell wall protuberances were discovered in the hypodermis at the zone of side root formation. These transfer cells were found neither in roots from the 0.5 mM NaCl treatment nor in the controls. Their possible role in the Na⁺/K⁺ selectivity of the roots of Prosopis farcta is discussed.     

不同耐盐性药用甘草幼苗根对Na~+的响应及其维管组织变化

以甘草属2种耐盐植物胀果甘草、乌拉尔甘草为材料,用不同浓度(50、100、150、200、250mmol·L-1)NaCl处理幼苗21d后,分析其生物量和根、茎、叶中的Na+、K+含量以及K+/Na+,计算根的离子选择吸收和运输系数,并应用光学显微镜观察比较二者的维管组织结构变化,以揭示2种药用甘草幼苗根对Na+的响应及其维管组织结构的变化特征,探讨甘草的耐盐机理。结果表明:(1)NaCl胁迫使2种甘草幼苗生物量均下降,在NaCl浓度为250mmol·L-1时,胀果甘草、乌拉尔甘草幼苗生物量分别是对照的53.34%、46.21%,胀果甘草耐盐性强于乌拉尔甘草。(2)随着NaCl浓度上升,2种甘草根积累的Na+显著增多,其中胀果甘草在所有盐处理下,根Na+含量均高于其它器官,说明其根对吸收的Na+具有显著截留效应;而乌拉尔甘草只在0~150mmol·L-1 NaCl范围内,根Na+含量显著高于叶片,当NaCl为200、250mmol·L-1时,叶片Na+含量显著高于根,说明乌拉尔甘草根对Na+的截留能力有限。(3)在相同盐处理下,胀果甘草离子选择吸收系数SAK,Na、离子运输系数STK,Na均大于乌拉尔甘草,胀果甘草根抑制Na+、促进K+向地上部运输的能力强于乌拉尔甘草。(4)乌拉尔甘草在NaCl为150、200mmol·L-1、胀果甘草在250mmol·L-1时,根结构对盐胁迫产生应激性响应,维管组织比例显著上升,有助于提高根向上的运输能力,减少盐害。研究表明,2种药用甘草根对Na+截留作用和向上运输时促K+抑Na+能力的差异,是导致其耐盐能力不同的主要原因,根对Na+的积累和截留作用的差异与根的结构响应相吻合,能较好地解释二者的耐盐性差异。     

NaCl胁迫对流苏幼苗生长、钠钾离子分布及渗透调节物质的影响

以2年生的流苏播种苗为材料,采用不同浓度(50、100、200、300 mmol·L^-1)NaCl溶液进行胁迫处理,研究盐胁迫对流苏的生长、Na^+和K^+分布格局、渗透调节物质的影响,以明确其耐盐阈值。结果表明:(1)随着NaCl胁迫浓度的增加,流苏幼苗生长量逐渐降低,盐害指数升高、存活率下降;幼苗耐盐阈值为98.693 mmol·L^-1(0.577%W/V)。(2)随着NaCl胁迫浓度的增加,流苏幼苗各器官中的Na^+含量持续增加,并在浓度为50 mmol·L^-1时表现为根>叶>茎,在其余各处理组表现为叶>根>茎;幼苗根、叶中的K^+含量表现为先增后减的变化趋势,茎中K^+含量总体表现为下降趋势,且器官中K^+含量表现为根>叶>茎;幼苗根部到茎部和茎部到叶部的离子选择性运输能力、各器官中的K^+/Na^+比值均呈下降趋势。(3)随着NaCl浓度的增加,流苏幼苗叶片可溶性糖、可溶性蛋白含量总体呈上升趋势,其脯氨酸含量呈先上升后下降的趋势。研究发现,流苏幼苗根系可通过对Na^+的吸收和累积来阻止其向地上部运输进而避免盐害发生;叶片和茎中通过提高对K^+的选择性吸收和累积,从而增大K^+/Na^+比值以减缓盐分对其生理代谢的伤害。     

The involvement of wheat U‐box E3 ubiquitin ligase TaPUB1 in salt stress tolerance

U‐box E3 ubiquitin ligases play important roles in the ubiquitin/26S proteasome machinery and in abiotic stress responses. TaPUB1‐overexpressing wheat (Triticum aestivum L.) were generated to evaluate its function in salt tolerance. These plants had more salt stress tolerance during seedling and flowering stages, whereas the TaPUB1‐RNA interference (RNAi)‐mediated knock‐down transgenic wheat showed more salt stress sensitivity than the wild type (WT). TaPUB1 overexpression upregulated the expression of genes related to ion channels and increased the net root Na⁺ efflux, but decreased the net K⁺ efflux and H⁺ influx, thereby maintaining a low cytosolic Na⁺/K⁺ ratio, compared with the WT. However, RNAi‐mediated knock‐down plants showed the opposite response to salt stress. TaPUB1 could induce the expression of some genes that improved the antioxidant capacity of plants under salt stress. TaPUB1 also interacted with TaMP (Triticum aestivum α‐mannosidase protein), a regulator playing an important role in salt response in yeast and in plants. Thus, low cytosolic Na⁺/K⁺ ratios and better antioxidant enzyme activities could be maintained in wheat with overexpression of TaPUB1 under salt stress. Therefore, we conclude that the U‐box E3 ubiquitin ligase TaPUB1 positively regulates salt stress tolerance in wheat.     

The heterotrimeric G‐protein β subunit,AGB1, plays multiple roles in the Arabidopsis salinity response

Salinity stress includes both osmotic and ionic toxicity. Sodium homeostasis is influenced by Na⁺ uptake and extrusion, vacuolar Na⁺ compartmentation and root to shoot Na⁺ translocation via transpiration. The knockout mutant of the Arabidopsis heterotrimeric G‐protein Gβ subunit, agb1, is hypersensitive to salt, exhibiting a leaf bleaching phenotype. We show that AGB1 is mainly involved in the ionic toxicity component of salinity stress and plays roles in multiple processes of Na⁺ homeostasis. agb1 mutants accumulate more Na⁺ and less K⁺ in both shoots and roots of hydroponically grown plants, as measured by inductively coupled plasma atomic emission spectrometry. agb1 plants have higher root to shoot translocation rates of radiolabelled ²⁴Na⁺ under transpiring conditions, as a result of larger stomatal apertures and increased stomatal conductance. ²⁴Na⁺ tracer experiments also show that ²⁴Na⁺ uptake rates by excised roots of agb1 and wild type are initially equal, but that agb1 has higher net Na⁺ uptake at 90 min, implicating possible involvement of AGB1 in the regulation of Na⁺ efflux. Calcium alleviates the salt hypersensitivity of agb1 by reducing Na⁺ accumulation to below the toxicity threshold. Our results provide new insights into the regulatory pathways underlying plant responses to salinity stress, an important agricultural problem.     

ZxAKT1 is essential for K+ uptake and K+/Na+ homeostasis in the succulent xerophyte Zygophyllum xanthoxylum

The inward‐rectifying K⁺ channel AKT1 constitutes an important pathway for K⁺ acquisition in plant roots. In glycophytes, excessive accumulation of Na⁺ is accompanied by K⁺ deficiency under salt stress. However, in the succulent xerophyte Zygophyllum xanthoxylum, which exhibits excellent adaptability to adverse environments, K⁺ concentration remains at a relatively constant level despite increased levels of Na⁺ under salinity and drought conditions. In this study, the contribution of ZxAKT1 to maintaining K⁺ and Na⁺ homeostasis in Z. xanthoxylum was investigated. Expression of ZxAKT1 rescued the K⁺‐uptake‐defective phenotype of yeast strain CY162, suppressed the salt‐sensitive phenotype of yeast strain G19, and complemented the low‐K⁺‐sensitive phenotype of Arabidopsis akt1 mutant, indicating that ZxAKT1 functions as an inward‐rectifying K⁺ channel. ZxAKT1 was predominantly expressed in roots, and was induced under high concentrations of either KCl or NaCl. By using RNA interference technique, we found that ZxAKT1‐silenced plants exhibited stunted growth compared to wild‐type Z. xanthoxylum. Further experiments showed that ZxAKT1‐silenced plants exhibited a significant decline in net uptake of K⁺ and Na⁺, resulting in decreased concentrations of K⁺ and Na⁺, as compared to wild‐type Z. xanthoxylum grown under 50 mm NaCl. Compared with wild‐type, the expression levels of genes encoding several transporters/channels related to K⁺/Na⁺ homeostasis, including ZxSKOR, ZxNHX, ZxSOS1 and ZxHKT1;1, were reduced in various tissues of a ZxAKT1‐silenced line. These findings suggest that ZxAKT1 not only plays a crucial role in K⁺ uptake but also functions in modulating Na⁺ uptake and transport systems in Z. xanthoxylum, thereby affecting its normal growth.     

Two closely linked tomato HKT coding genes are positional candidates for the major tomato QTL involved in Na+/K+ homeostasis

The location of major quantitative trait loci (QTL) contributing to stem and leaf [Na⁺] and [K⁺] was previously reported in chromosome 7 using two connected populations of recombinant inbred lines (RILs) of tomato. HKT1;1 and HKT1;2, two tomato Na⁺‐selective class I‐HKT transporters, were found to be closely linked, where the maximum logarithm of odds (LOD) score for these QTLs located. When a chromosome 7 linkage map based on 278 single‐nucleotide polymorphisms (SNPs) was used, the maximum LOD score position was only 35 kb from HKT1;1 and HKT1;2. Their expression patterns and phenotypic effects were further investigated in two near‐isogenic lines (NILs): 157‐14 (double homozygote for the cheesmaniae alleles) and 157‐17 (double homozygote for the lycopersicum alleles). The expression pattern for the HKT1;1 and HKT1;2 alleles was complex, possibly because of differences in their promoter sequences. High salinity had very little effect on root dry and fresh weight and consequently on the plant dry weight of NIL 157‐14 in comparison with 157‐17. A significant difference between NILs was also found for [K⁺] and the [Na⁺]/[K⁺] ratio in leaf and stem but not for [Na⁺] arising a disagreement with the corresponding RIL population. Their association with leaf [Na⁺] and salt tolerance in tomato is also discussed.     

盐胁迫对3种平欧杂种榛幼苗叶片解剖结构及离子吸收、运输与分配的影响

研究盐胁迫下3个品种平欧杂种榛幼苗叶片解剖结构和离子代谢特征,以揭示盐胁迫响应与适应机制及不同品种的耐盐性差异。以‘达维’、‘辽榛7号’、‘玉坠’2年生压条苗为材料,在盆栽条件下经轻度、中度、重度(分别为50、100、200 mmol/L NaCl)盐胁迫处理,设对照为0,研究幼苗叶片显微解剖结构参数和Na~+、K~+、Cl~-、Ca²⁺含量的变化及其在根、茎、叶中的吸收、运输和分配特征。不同品种平欧杂种榛叶片厚度、上表皮厚度、下表皮厚度、栅栏组织和海绵组织厚度随着盐胁迫程度的增强呈现出先增加后降低的特点,轻度和中度胁迫下各参数显著高于对照。中度盐胁迫显著提高了各品种叶片结构紧密度。盐胁迫导致平欧杂种榛根、茎、叶Na~+和Cl~-含量明显高于对照。盐胁迫下,Na~+和Cl~-在叶中的绝对含量明显高于茎和根,但二者的增幅以根中最大,叶中最小,表明平欧杂种榛根系首先会吸收并截留一定数量的Na~+和Cl~-,然后将其运输至茎和叶中。与对照相比,轻度和中度盐胁迫下根、茎对K~+和Ca²⁺的吸收保持稳定或减少,叶对K~+和Ca²⁺...     

Differences in efficient metabolite management and nutrient metabolic regulation between wild and cultivated barley grown at high salinity

Physiological and biochemical responses of Hordeum maritimum and H. vulgare to salt stress were studied over a 60‐h period. Growth at increasing salinity levels (0, 100, 200 and 300 mM NaCl) was assessed in hydroponic culture. H. maritimum was shown to be a true halophyte via its typical behaviour at high salinity. Shoot growth of cultivated barley was gradually reduced with increasing salinity, whereas that of wild barley was enhanced at 100 and 200 mm NaCl then slightly reduced at 300 mM NaCl. The higher salt tolerance of H. maritimum as compared to H. vulgare was due to its higher capacity to maintain cell turgor under severe salinity. Furthermore, H. maritimum exhibited fine regulation of Na⁺ transport from roots to shoots and, unlike H. vulgare, it accumulated less Na⁺ in shoots than in roots. In addition, H. maritimum can accumulate more Na⁺ than K⁺ in both roots and shoots without the appearance of toxicity symptoms, indicating that Na⁺ was well compartmentalized within cells and substituted K⁺ in osmotic adjustment. The higher degree of salt tolerance of H. maritimum is further demonstrated by its economic strategy: at moderate salt treatment (100 mm NaCl), it used inorganic solutes (such as Na⁺) for osmotic adjustment and kept organic solutes and a large part of the K⁺ for metabolic activities. Indeed, K⁺ use efficiency in H. maritimum was about twofold that in H. vulgare; the former started to use organic solutes as osmotica only at high salinity (200 and 300 mm NaCl). These results suggest that the differences in salt tolerance between H. maritimum and H. vulgare are partly due to (i) differences in control of Na⁺ transport from roots to shoots, and (ii) H. maritimum uses Na⁺ as an osmoticum instead of K⁺ and organic solutes. These factors are differently reflected in growth.     

Arbuscular mycorrhizal fungi native from a Mediterranean saline area enhance maize tolerance to salinity through improved ion homeostasis

Soil salinity restricts plant growth and productivity. Na⁺ represents the major ion causing toxicity because it competes with K⁺ for binding sites at the plasma membrane. Inoculation with arbuscular mycorrhizal fungi (AMF) can alleviate salt stress in the host plant through several mechanisms. These may include ion selection during the fungal uptake of nutrients from the soil or during transfer to the host plant. AM benefits could be enhanced when native AMF isolates are used. Thus, we investigated whether native AMF isolated from an area with problems of salinity and desertification can help maize plants to overcome the negative effects of salinity stress better than non‐AM plants or plants inoculated with non‐native AMF. Results showed that plants inoculated with two out the three native AMF had the highest shoot dry biomass at all salinity levels. Plants inoculated with the three native AMF showed significant increase of K⁺ and reduced Na⁺ accumulation as compared to non‐mycorrhizal plants, concomitantly with higher K⁺/Na⁺ ratios in their tissues. For the first time, these effects have been correlated with regulation of ZmAKT2, ZmSOS1 and ZmSKOR genes expression in the roots of maize, contributing to K⁺ and Na⁺ homeostasis in plants colonized by native AMF.     

Heterologous expression of Arabidopsis H+‐pyrophosphatase enhances salt tolerance in transgenic creeping bentgrass (Agrostis stolonifera L.)

The Arabidopsis vacuolar H⁺‐pyrophosphatase (AVP1), when over‐expressed in transgenic (TG) plants, regulates root and shoot development via facilitation of auxin flux, and enhances plant resistance to salt and drought stresses. Here, we report that TG perennial creeping bentgrass plants over‐expressing AVP1 exhibited improved resistance to salinity than wild‐type (WT) controls. Compared to WT plants, TGs grew well in the presence of 100 mm NaCl, and exhibited higher tolerance and faster recovery from damages from exposure to 200 and 300 mm NaCl. The improved performance of the TG plants was associated with higher relative water content (RWC), higher Na⁺ uptake and lower solute leakage in leaf tissues, and with higher concentrations of Na⁺, K⁺, Cl^‐ and total phosphorus in root tissues. Under salt stress, proline content was increased in both WT and TG plants, but more significantly in TGs. Moreover, TG plants exhibited greater biomass production than WT controls under both normal and elevated salinity conditions. When subjected to salt stress, fresh (FW) and dry weights (DW) of both leaves and roots decreased more significantly in WT than in TG plants. Our results demonstrated the great potential of genetic manipulation of vacuolar H⁺‐pyrophosphatase expression in TG perennial species for improvement of plant abiotic stress resistance.     

Contributions of arbuscular mycorrhizal fungi to growth,photosynthesis, root morphology and ionic balance of citrus seedlings under salt stress

A pot study was conducted to determine the effects of arbuscular mycorrhizal (AM) fungi (Glomus mosseae and Paraglomus occultum) and salt (NaCl) stress on growth, photosynthesis, root morphology and ionic balance of citrus (Citrus tangerine Hort. ex Tanaka) seedlings. Eighty-five-day-old seedlings were exposed to 100 mM NaCl for 60 days to induce salt stress. Mycorrhizal colonization of citrus seedlings was not affected by salinity when associated with P. occultum, but significantly decreased when with G. mosseae. Compared with the non-mycorrhizal controls, mycorrhizal seedlings generally had greater plant height, stem diameter, shoot, root and total plant biomass, photosynthetic rate, transpiration rate and stomatal conductance under the 0 and 100 mM NaCl stresses. Root length, root projected area and root surface area were also higher in the mycorrhizal than in the non-mycorrhizal seedlings, but higher root volume in seedlings with G. mosseae. Leaf Na⁺ concentrations were significantly decreased, but leaf K⁺ and Mg²⁺ concentrations and the K⁺/Na⁺ ratio were increased when seedlings with both G. mosseae and P. occultum. Under the salt stress, Na⁺ concentrations were increased but K⁺ concentrations decreased in the mycorrhizal seedlings. Under the salt stress, Ca²⁺ concentrations were increased in the seedlings with P. occultum or without AM fungi (AMF), but decreased with G. mosseae. Ratios of both Ca²⁺/Na⁺ and Mg²⁺/Na⁺ were also increased in seedlings with G. mosseae under the non-salinity stress, while only the Mg²⁺/Na⁺ ratio was increased in seedlings with P. occultum under the salt stress. Our results suggested that salt tolerance of citrus seedlings could be enhanced by associated AMF with better plant growth, root morphology, photosynthesis and ionic balance.     

SbHKT1;4, a member of the high-affinity potassium transporter gene family from Sorghum bicolor,functions to maintain optimal Na＋/K＋ balance under Na＋ stress

In halophytic plants, the high-affinity potassium transporter HKT gene family can selectively uptake K＋ in the presence of toxic concentrations of Na＋. This has so far not been well examined in glycophytic crops. Here, we report the characterization of SbHKTI;4, a member of the HKT gene family from Sorghum bicolor. Upon Na＋ stress, SbHKT1;4 expression was more strongly upregulated in salt-tolerant sorghum accession, correlating with a better balanced Na＋/ K＋ ratio and enhanced plant growth. Heterogeneous expression analyses in mutants of Saccharomyces cerevisiae and Arabidopsis thaliana indicated that overexpressing SbHKT1;4 resulted in hypersensitivity to Na＋ stress, and such hypersensitivity could be alleviated with the supply of elevated levels of K＋, implicating that SbHKT1;4 may mediate K＋ uptake in the presence of excessive Na＋. Further electrophysiological evidence demonstrated that SbHKT1;4 could transport Na＋ and K＋ when expressed in Xenopus laevis oocytes. The relevance of the finding that SbHKTI;4 functions to maintain optimal Na＋/K＋ balance under Na＋ stress to the breeding of salt-tolerant glycophytic crops is discussed.     

Interactions among Ca2+, Na+ and K+ in salinity toxicity: quantitative resolution of multiple toxic and ameliorative effects

Root elongation by wheat seedlings (Triticum aestivum L. cv. Scout 66) was not inhibited by NaCl or KCl up to 130 mM in culture solutions or by high Na⁺ (2 mg g^-1 FW) or K⁺ (4 mg g^-1 FW) in the root tissue, provided that [Ca²⁺]>2 mM in the rooting medium. At [NaCl], [KCl], or [mannitol] >250 mOs, root elongation was progressively inhibited, irrespective of high [Ca²⁺]. In contrast, shoot elongation was sensitive to any diminution of water potential, and Ca²⁺ alleviated the toxicity only weakly. At solute concentrations <250 mOs, the following interactions were observed. Ca²⁺ alleviated Na⁺ and K⁺ toxicity to roots by at least three separate mechanisms. K⁺ was more toxic to roots than Na⁺, but Na⁺ was more toxic to shoots. Low levels of K⁺ relieved Na⁺ toxicity, but low levels of Na⁺ enhanced K⁺ toxicity. Tissue concentrations of Na⁺ were reduced by Ca²⁺ and K⁺ in the rooting medium, and tissue concentrations of K⁺ were enhanced by Ca²⁺ and Na⁺. Several hypotheses relating to salinity toxicity can be evaluated, at least for wheat seedlings. The osmoticant hypotheses (salinity intoxication occurs because of diminished water potential) is true for shoots at all salinity levels, but is true for roots only at high salinity. The Ca²⁺-displacement hypothesis (Na⁺ is toxic because it displaced Ca²⁺ from the cell surface) is correct, but often of minor importance. The K⁺-depletion hypothesis (Na⁺ is toxic because it causes a loss of K⁺ from plant tissues) is false. The Cl^--toxicity hypothesis (the apparent toxicity of Na⁺ is induced by associated Cl^-) is false. The results indicate that, apart from osmotic effects, high levels of Na⁺ in the rooting medium and in the tissues are not toxic unless Ca²⁺ is also deficient, a condition probably leading to inadequate compartmentation and excessive cytoplasmic accumulation. This study related growth to ion activities at plasma-membrane surfaces. These activities were computed by a Gouy-Chapman-Stern model then incorporated into non-linear growth models for growth versus toxicants and ameliorants.Key words: Calcium, potassium, salinity, sodium, toxicity