Some mechanisms of salt tolerance in crop plants

Summary In the first part of this review the main features of salt tolerance in higher plants are discussed. The hypothesis of intracellular compartmentation of solutes is used as a basis for models of tolerance mechanisms operating in roots and in leaves. Consideration is given to the implications of the various mechanisms for the yield potential of salt-tolerant crop plants.Some work on the more salt-tolerant members of the Triticeae is then described. The perennial speciesElytrigia juncea andLeymus sabulosus survive prolonged exposure to 250 mol m^–3 NaCl, whereas the annual Triticum species are severely affected at only 100 mol m^–3 NaCl. In the perennial species the tissue ion levels are controlled within narrow limits. In contrast, the more susceptible wheats accumulate far more sodium and chloride than is needed for osmotic adjustment, and the effects of salt stress increase with time of exposure.Two different types of salt tolerance are exhibited in plants capable of growing at high salinities. In succulent Chenopodiaceae, for example, osmotic adjustment is achieved mainly by accumulation of high levels of sodium and chloride in the shoots, accompanied by synthesis of substantial amounts of the compatible solute glycinebetaine. This combination of mechanisms allows high growth rates, in terms of both fresh and dry weight. At the opposite end of the spectrum of salt tolerance responses are the halophytic grasses, which strictly limit the influx of salts into the shoots, but suffer from very much reduced growth rates under saline conditions. Another variation is shown in those species that possess salt glands. The development and exploitation of crop plants for use on saline soils is discussed in relation to the implications of these various mechanisms.     

Selection of callus cultures of sugarcane (Saccharum sp.) tolerant to NaCl and their response to salt stress

Stable callus cultures tolerant to NaCl (68 mM) were developed from salt-sensitive sugarcane cultivar CP65-357 by in vitro selection process. The accumulation of both inorganic (Na⁺, Cl⁻ and K⁺) and organic (proline and soluble sugars) solutes was determined in selected and non-selected calli after a NaCl shock in order to evaluate their implication in in vitro salt tolerance of the selected lines. Both salt-tolerant and non-selected calli showed similar relative fresh weight growth in the absence of NaCl. No growth reduction was observed in salt-tolerant calli while a significant reduction about 32% was observed in nonselected ones when both were cultivated on 68 mM NaCl. Accumulation of Na⁺ was similar in both salt-tolerant and non-selected calli in the presence of NaCl. Accumulation of Cl⁻ was lower in NaCl-tolerant than in non-selected calli while proline and soluble sugars were more accumulated in salt-tolerant than in non-selected calli when both were exposed to salt. K⁺ level decreased more severely in non-selected calli than in NaCl-tolerant ones after NaCl shock. The results indicated that K⁺ and Cl⁻ may play a key role in in vitro salt-tolerance in sugarcance cell lines obtained by in vitro selection and that organic solutes could contribute mainly to counteract the negative water potential of the outside medium.     

Physiological and molecular aspects of salt stress in plants

The study of salt stress mechanisms in plants has become an important issue for the modern agricultural development, climate change, and global food crisis. The plant response to high salt concentrations is complex and comprehensive; it includes many different processes, which should be correctly coordinated. The effect of excessive salt concentrations on plants results in osmotic stress and creates an ionic inbalance due to the accumulation of toxic ions, such as Cl^? and, especially, Na⁺. Salt stress also has negative impact on mineral homeostasis, in particular Ca²⁺ and K⁺. The progress in transcryptomics, genomics, and molecular biology revealed a new gene families that participate in the formation of salt stress response in plants. This review describes the fundamental principles and mechanisms of plant salt tolerance, maintenance of ion homeostasis. In this paper the detailed analysis of the maine transport membrane systems responsible for the transport of ions and their role in plant salt stress were conducted. The perspectives and directions for the further biotechnological and genetic improvement of salt tolerance in plants are underlied.     

藜异型性种子后代植株盐响应生理机制

种子异型性是植物适应异质生境的重要策略,异型性对后代植株的影响值得深入研究。以具有种子异型性的荒漠植物藜为材料,测定了异型种子后代植株在不同浓度(0,50,300 mmol/L)Na Cl胁迫下的表型及与耐盐相关的生理指标变化,探讨了异型种子后代植株之间对盐胁迫生理响应机制的差异。结果显示,褐色种子后代植株在各浓度盐胁迫下的表型(株高、分枝数)均优于黑色种子。对其生理指标测定结果显示,褐色种子植株除电导率显著低于黑色种子植株外,氧化损伤指标(氧自由基、丙二醛含量)及抗氧化酶(SOD、CAT、POX)活性、抗氧化剂(Car、As A)含量、渗调物质(可溶性糖、脯氨酸、甜菜碱)含量等在两者之间无明显差异。研究结果暗示,藜异型性种子后代植株的早期生长表型在盐胁迫下所产生的差异可能是由种子胚的大小及萌发快慢差异所导致,而这种差异并未引起盐胁迫下两种植株抗氧化系统响应的差异。     

甘薯愈伤组织对干旱胁迫和盐胁迫的生理反应对比

研究干旱胁迫和盐胁迫对“芦选一号”。日‘薯愈伤组织可溶性蛋白、可溶性糖、脯氨酸含量、SOD活性等的影响,从而在细胞水平上探讨甘薯抵御渗透胁迫的生理机制。并分析甘薯细胞对干旱处理（PEG-6000）和盐处理（NaCl）的反应差异。结果表明,可溶性蛋白质含量在干旱胁迫下缓慢升高,在轻度和中度盐胁迫的生长前期和中期有较大幅度的上升。但后期下降,表明短时间盐胁迫下,Na^＋可能促进可溶性蛋白的合成;MDA在重度干旱胁迫下的含量显著低于重度盐胁迫,而SOD活性显著高于盐胁迫。表明在盐胁迫下细胞膜透性增加的主要原网是膜脂过氧化作用。干旱处理则是PEG-6000脱水的直接结果;重度干旱胁迫下,可溶性糖含量在短期内迅速升高,然后下降,而脯氨酸含量则在胁迫中后期迅速上升。脯氨酸可能有补偿可溶性糖含量降低的作用。     

Physiological responses of four herbaceous plants to aluminum stress in South China

Different plants have physiological responses under A1 stress,but there is no systematic study to examine physiological responses of herbaceous plants under Al stress.The aim of this study is to investigate the effect of Al on physiological characteristics of four herbaceous plants,which distributed in red soil area in South China,and to analyze the differences in physiological responses to Al stress between the four herbaceous plants.Four herbaceous plants(Pharbitis nil,Cassia occidentlis,Echinochloa colonum and Aeschynomene indica)were used,and the seed germination percentage,the contents of chlorophyll,proline,and malondialdehyde(MDA),membrane permeability(MP),soluble sugar,and activities of peroxides(POD)and cata lase(CAT)in leaves under five Al3+ treatments(0,80,400,2000,and 10000 mg/L)were assayed with the sand culture method.The results showed remarkable effects of Al3+ on physiological characteristics of these four herbaceous plants.The seeds of all the four species could not germinate at 10000 mg/L.and the growth of all plants were retarded under the 2000 mg/LAr3+ treatment.Compared with the control,2000 mg/L Al3+ significantly(P＜0.05)reduced the contents of chlorophyll a and chlorophyll a+b,and increased the contents of MDA and MP.The content of proline in creased very significantly(P＜0.01)and activities of POD and CAT were depressed.The contents of MDA and MP in leaves of P.nil and A.indica decreased,and the activities of POD and CAT in leaves of the two plants increased under 80mg/L and 400 mg/L.However,the changes in C.occidentlis leaves were opposite to those of the above two plants.The changes in leaves of E.colonum were similar to those of P.nil and A.indica at 80 mg/L.but were opposite to those at 400mg/LAr3+.It is suggested that plants with higher activities of POD and CAT,more contents of chlorophyll and proline,and lower contents of MDA and MP consequently improve the tolerance to Al stress under low and middle Al treatments.     

Physiological responses of four herbaceous plants to aluminum stress in South China

Different plants have physiological responses under Al stress, but there is no systematic study to examine physiological responses of herbaceous plants under Al stress. The aim of this study is to investigate the effect of Al on physiological characteristics of four herbaceous plants, which distributed in red soil area in South China, and to analyze the differences in physiological responses to Al stress between the four herbaceous plants. Four herbaceous plants (Pharbitis nil, Cassia occidentlis, Echinochloa colonum and Aeschynomene indica) were used, and the seed germination percentage, the contents of chlorophyll, proline, and malondialdehyde (MDA), membrane permeability (MP), soluble sugar, and activities of peroxides (POD) and catalase (CAT) in leaves under five Al³⁺ treatments (0, 80, 400, 2 000, and 10 000 mg/L) were assayed with the sand culture method. The results showed remarkable effects of Al³⁺ on physiological characteristics of these four herbaceous plants. The seeds of all the four species could not germinate at 10 000 mg/L, and the growth of all plants were retarded under the 2 000 mg/L Al³⁺ treatment. Compared with the control, 2 000 mg/L Al³⁺ significantly (P < 0.05) reduced the contents of chlorophyll a and chlorophyll a + b, and increased the contents of MDA and MP. The content of proline increased very significantly (P < 0.01) and activities of POD and CAT were depressed. The contents of MDA and MP in leaves of P. nil and A. indica decreased, and the activities of POD and CAT in leaves of the two plants increased under 80 mg/L and 400 mg/L. However, the changes in C. occidentlis leaves were opposite to those of the above two plants. The changes in leaves of E. colonum were similar to those of P. nil and A. indica at 80 mg/L, but were opposite to those at 400 mg/L Al³⁺. It is suggested that plants with higher activities of POD and CAT, more contents of chlorophyll and proline, and lower contents of MDA and MP consequently improve the tolerance to Al stress under low and middle Al treatments. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(4): 644–651 [译自: 植物生态学报, 2005, 29(4): 644–651]     

Physiological responses to water stress following a conditioning period in berseem clover

Berseem clover (Trifolium alexandrinum L.) is an important crop in semi-arid regions; its herbage and seed yields are often reduced by water stress. Our objectives were (i) to determine the effect of water stress, applied after a conditioning period, on water relations, proline accumulation and plant dry weight, and (ii) to investigate if some physiological responses differed in varieties of berseem. Five cultivars (Axi, Bigbee, Lilibeo, Sacromonte and Saniros) were grown in a controlled environment, and subjected to four irrigation treatments (T1, T2, T3 and T4 referring to plants irrigated to field capacity every 1, 2, 3 or 4 d, respectively) during a conditioning period (12 d). T1 treatment indicated the well-watered control, whereas T2, T3 and T4 treatments represented the conditioned plants. Leaf water potential (Ψ), osmotic potential (Ψπ), relative water content (RWC), gravimetric soil water content (GSWC) and leaf proline concentration were recorded during the conditioning period and a subsequent water deficit period (3 d) applied at early flowering growth stage. The conditioned plants subjected to subsequent water deficit maintained higher values of Ψ, Ψπ, RWC and GSWC, and lower values of leaf proline concentration. Reductions in parameter values were inversely related to the water stress severity that plants had previously experienced. At the end of the experiment, T1 showed 42%, 58% and 31% lower values for Ψ, Ψπ and RWC, respectively, than those of T4. Conditioned plants were also shorter and accumulated less leaf, stem and total dry weight. The conditioning treatments did not affect the relation between Ψ and Ψπ since conditioned plants show similar values of Ψπ as the control at the same Ψ value. Thus, drought acclimation in berseem clover contributed to water stress tolerance by the maintenance of tissue hydration. The berseem cultivars examined showed differences in plant growth parameters, but they were very similar for physiological responses to water deficit. The main genetic difference was recorded for turgor maintenance capacity. This revised version was published online in June 2006 with corrections to the Cover Date.     

Na+ accumulation in root symplast of sunflower plants exposed to moderate salinity is transpiration-dependent

Twenty-day-old sunflower plants (Helianthus annuus L. cv Sun-Gro 380) grown hydroponically under controlled conditions were used to study the effect of transpiration on Na(+) compartmentalization in roots. The plants were exposed to low Na(+) concentrations (25mM NaCl) and different environmental humidity conditions over a short time period (8.5h). Under these conditions, Na(+) was accumulated primarily in the root, but only the Na(+) accumulated in the root symplast was dependent on transpiration, while the Na(+) accumulated in both the shoot and the root apoplast exhibited a low transpiration dependence. Moreover, Na(+) content in the root apoplast was reached quickly (0.25h) and increased little with time. These results suggest that, in sunflower plants under moderate salinity conditions, Na(+) uptake in the root symplast is mediated by a transport system whose activity is enhanced by transpiration.     

两种野生灌木幼苗对干旱和盐交叉胁迫的生理响应

长枝木蓼(Ateaphaxis viegata)和刺叶锦鸡儿(Caragana acanthophulla)是乌鲁木齐周边植被组成的重要种,在植被恢复中具有潜在价值。该文通过盆栽控水控盐法研究两种野生灌木幼苗在不同程度的干旱和盐交叉胁迫下的生理反应。结果表明,干旱和盐交叉胁迫下,长枝木蓼可溶性糖增幅较刺叶锦鸡儿大;轻度交叉胁迫时,两种灌木的可溶性糖含量高于仅干旱胁迫或盐胁迫下的。中度和重度交叉胁迫下,长枝木蓼叶片丙二醛含量增幅高于刺叶锦鸡儿。交叉胁迫下,两种灌木叶片叶绿素含量下降,且长枝木蓼叶片的叶绿素含量降幅大于刺叶锦鸡儿。因膜系统的过氧化作用,使MDA含量升高。轻度干旱胁迫提高了两种野生灌木对盐胁迫的耐受能力。两种灌木能很好地适应中度交叉胁迫,重度交叉胁迫对两种灌木的伤害很大,但也能生存。综合比较各参数发现,刺叶锦鸡儿更耐早,而长枝木蓼更耐盐;综合比较各生理参数发现,刺叶锦鸡儿对干旱和盐胁迫的耐受能力较长枝木蓼强。     

盐胁迫下根瘤共生豌豆植株对外源钙的生理响应

为明确外源钙对根瘤共生豌豆植株耐盐性的影响机制,本试验采用盆栽方式研究了NaCl (170 mmol·L^-1)胁迫下,外源施加CaCl₂(0、5、15 mmol·L^-1)对接种根瘤菌(菌株15657、15735、Ca66)的两种耐盐性不同品种豌豆(定豌8号、陇豌6号)植株生理指标的影响。结果表明: 接种根瘤菌、施加CaCl₂或接种根瘤菌后施加CaCl₂均提高了豌豆植株的生物量、超氧化物歧化酶(SOD)和过氧化物酶(POD)活性、脯氨酸(Pro)和可溶性糖(SS)含量,降低了丙二醛(MDA)含量;接种根瘤菌后施加15 mmol·L^-1 CaCl₂显著提高了豌豆植株的生物量、POD活性和Pro含量。接种与豌豆植株匹配性较好的15735菌株并施加CaCl₂对豌豆在盐胁迫下各指标的影响相对较小,接种匹配性差的菌株(156567、Ca66)并施加CaCl₂影响较大。经隶属函数综合分析表明,接种根瘤菌后施加CaCl₂的豌豆植株表现出较强的耐盐性,接种15735菌后施加15 mmol·L^-1 CaCl₂的定豌8号隶属函数值为0.814,耐盐性最强。本研究表明,相比接种根瘤菌和施加CaCl₂处理,接种根瘤菌后施加CaCl₂可更有效地提高盐胁迫下豌豆植株的抗氧化酶活性,增强渗透调节能力,降低膜脂过氧化伤害,从而提高豌豆植株的耐盐性。     

Physiological and ultrastructural responses of Catharanthus roseus cell suspension to salt stress

Catharanthus roseus (L.) cell response to salinity was investigated. Seven days after cell treatment with 100 mM NaCl, they showed a decrease in dry weight and an increase in sodium and chloride contents (about 12.4- and 1.5-fold, respectively, in comparison to the control). At the ultrastructural level, NaCl treatment reduced cell size and increased plastid density. In addition, it reduced the starch grain size and their number per plastid; however, starch content was 1.5-fold increased, which was due to the increase in the plastid density. At the ultrastructural level, the applied salinity had no obvious effects, such as swelling or disorganization of plastids except a slight decrease in the stroma electron density. Equally, no deleterious effect was observed on mitochondria except a small increase of their crista volume and matrix electron density. It was shown that, although the relative sensitivity of C. roseus cells to salt stress pointed by the reduction in the dry weight, a decrease in the cell size, and the high accumulation of toxic ions, they preserved the integrity of their plastids and mitochondria.     

Nitrogen metabolism of young barley plants as affected by NaCl-salinity and potassium

Summary In a solution culture experiment with 31 days old barley plants (var. Miura) the influence of NaCl-salinization (80 mM) and KCl addition (5 and 10 mM) on the uptake and turnover of labelled nitrogen (¹⁵NH₄ ¹⁵NO₃) was studied. Labelled N was applied for 24 h at the end of a 20 days' salinization period. Salinization impaired growth and incorporation of labelled N into the protein fraction paralleled by accumulation of labelle dinorganic N. All salt effects were much more pronounced in the shoots than in the roots.Potassium addition enhanced N uptake (total¹⁵N-content) and incorporation into protien, reduced the accumulation of inorganic N and improved the growth of salinized plants.The presented data support the point of view that impairment of protein (enzyme) metabolism is an important aspect of salt stress which is probably induced by the disturbance of the K/Na balance of the tissues under saline conditions.This work was supported by a grant from the Alexander von Humboldt foundation.     

Physiological responses of Pinus sylvestris to atmospheric ammonia

Summary Young saplings of Pinus sylvestris were fumigated for 3 months with ammonia in concentrations ranging from 0 to 240 g m^-3. Despite the much higher concentrations than normal in the field, no visible damage occurred. Photosynthesis, dark respiration, transpiration and biomass production were stimulated. At 240 g m^-3 with high irradiance (PAR: 950 mol m^-2 s^-1), net photosynthesis was stimulated by 24% and dark respiration by 76%. Intitial light use efficiency was not significantly affected. Transpiration increased, both in the dark and at 950 mol m^-2 s^-1 by 40% and 57%, respectively. In the presence of ammonia, stomatal control was less efficient. Though growth of roots was not affected by NH₃, that of current year needles was stimulated, resulting in an increased mass ratio of needles to roots. The nitrogen content of the needles increased, but the contents of other mineral components did not change significantly. Due to increased transpiration per unit of needle area and increased mass of needles per tree, water loss per tree was about twice as high in the treatment with 240 g m^-3 as in the control. Towards the end of fumigation, a 10-day period without water supply followed and then the water potential of the shoots was measured as an indicator of water demand. This demand was higher with higher concentrations of NH₃, suggesting a higher risk of injury from drought.     

Short-term and long-term effects of plant water deficits on stomatal response to humidity in Corylus avellana L.

Short-term (hours) changes in plant water status were induced in hazel (Corylus avellana L.) by changing the evaporative demand on a major portion of the shoot while maintaining a branch in a constant environment. Stomatal conductance of leaves on the branch was influenced little by these short-term changes in water status even with changes in leaf water potential as great as 8 bars. Long-term (days) changes in plant water status were imposed by soil drying cycles. Stomatal conductance progessively decreased with increases in long-term water stress. Stomata still responded to humidity with long-term water stress but the range of the conductance response decreased. Threshold responses of stomata to leaf water potential were not observed with either short-term or long-term changes in plant water status even when leaves wilted. It is suggested that concurrent measurements of plant water status may not be sufficient for explaining stomatal and other plant responses to drought.     

Dynamic metabonomic responses of tobacco (Nicotiana tabacum) plants to salt stress

Metabolic responses are important for plant adaptation to osmotic stresses. To understand the dosage and duration dependence of salinity effects on plant metabolisms, we analyzed the metabonome of tobacco plants and its dynamic responses to salt treatments using NMR spectroscopy in combination with multivariate data analysis. Our results showed that the tobacco metabonome was dominated by 40 metabolites including organic acids/bases, amino acids, carbohydrates and choline, pyrimidine, and purine metabolites. A dynamic trajectory was clearly observable for the tobacco metabonomic responses to the dosage of salinity. Short-term low-dose salt stress (50 mM NaCl, 1 day) caused metabolic shifts toward gluconeogenesis with depletion of pyrimidine and purine metabolites. Prolonged salinity with high-dose salt (500 mM NaCl) induced progressive accumulation of osmolytes, such as proline and myo-inositol, and changes in GABA shunt. Such treatments also promoted the shikimate-mediated secondary metabolisms with enhanced biosynthesis of aromatic amino acids. Therefore, salinity caused systems alterations in widespread metabolic networks involving transamination, TCA cycle, gluconeogenesis/glycolysis, glutamate-mediated proline biosynthesis, shikimate-mediated secondary metabolisms, and the metabolisms of choline, pyrimidine, and purine. These findings provided new insights for the tobacco metabolic adaptation to salinity and demonstrated the NMR-based metabonomics as a powerful approach for understanding the osmotic effects on plant biochemistry.     

Response of plants to CO2 under water limited conditions

The influence of inefeased atmospheric CO₂ on the interaction between plant growth and water use is proving to be one of the most profound impacts of the anthropogenic Greenhouse Effect. This paper illustrates the interaction between CO₂ and water in plant growth at a range of scales. Most published work has concentrated on water use efficiency, especially at shorter time scales, and has shown large increases of leaf water use efficiency with increased CO₂. However, the magnitude of the effect is variable, and does not consistently agree with predictions from simple leaf gas exchange considerations. The longer the time scales considered, the less the information and the more the uncertainty in the response to CO₂, because of the additional factors that have to be considered, such as changes in leaf area, respiration of non-photosynthetic tissues and soil evaporation. The need for more detailed studies of the interactions between plant evaporation, water supply, water status and growth is stressed, as increased CO₂ can affect all of these either directly, or indirectly through feedbacks with leaf gas exchange, carbon partitioning, leaf growth, canopy development and root growth.     

花期海蓬子对盐胁迫的生理响应

研究了花期阶段不同浓度NaCl(0、10‰、20‰、30‰、40‰、50‰和60‰)对海蓬子(Salicornia bigelovii)生长、光合色素、光合作用参数、抗氧化和离子含量的影响.结果表明:10‰NaCl盐处理下海蓬子株高、茎生物量、叶绿素(Chl)含量、净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)、胞间CO2浓度(Ci)、茎SOD活性和POD活性等指标值均显著高于其它处理,而高盐(40‰NaCl及其以上)胁迫下,上述指标值均表现为下降,同时茎K+含量也显著下降,而茎Na+、Cl、Na+/K+和MDA含量等显著上升.相关分析显示,生物量与Chl a/Chl b呈现显著正相关,与株高、茎Pn、Gs、Ci、Chl、Chl含量、K+含量、SOD和POD活性呈现极显著正相关,与气孔限制值(Ls)、Car/Chl、MDA含量、Cl含量和Na+/K+等均呈极显著负相关.综上所述,10‰NaCl处理是花期海蓬子生长和光合生长的最适宜盐度,而无盐和高盐下海蓬子光合抑制主要是来自气孔因素,同时高盐胁迫下还伴随着非气孔限制.