Physiological responses and adaptive strategies of wheat seedlings to salt and alkali stresses

Abstract

Effects of salt (NaCl?:?Na₂SO₄) and alkali (NaHCO₃?:?Na₂CO₃) stresses on the contents of inorganic ions and organic solutes in wheat shoots were compared to explore the physiological responses and adaptive strategies of wheat to these stresses. Wheat significantly accumulated Na⁺ and simultaneously accumulated Cl^?, soluble sugars and proline to maintain osmotic and ionic balance under salt stress. Compared with salt stress, the high pH from alkali stress enhanced Na⁺ accumulation and affected the absorption of inorganic anions. To maintain ionic and osmotic balance, wheat accumulated organic acids, soluble sugars and proline. The accumulation of Cl^? and organic acids was the main difference in the physiological responses and adaptive mechanisms to salt and alkali stresses, respectively.     

Effects of saline and alkaline stresses on the germination, growth, photosynthesis, ionic balance and anti-oxidant system in an alkali-tolerant leguminous forage Lathyrus quinquenervius

We compared the effects of saline stress (9:1 molar ratio of NaCl : Na₂SO₄, pH 6.44–6.65) and alkaline stress (9:1 molar ratio of NaHCO₃ : Na₂CO₃, pH 8.71–8.89) on the germination, growth, photosynthesis, ionic balance and activity of anti-oxidant enzymes of Lathyrus quinquenervius to elucidate the physiological adaptive mechanism of plants to alkaline stress (high pH). The results showed that, at a low stress intensity, the effects of saline stress and alkaline stress on L. quinquenervius were similar. Compared with saline stress, high alkaline stress intensity clearly inhibited germination, growth, photosynthesis and root system activity, and led to a sharp increase in Na⁺ and an ion imbalance in the shoots, as well as enhanced H₂O₂ and malondialdehyde content, resulting in severe intracellular oxidative stress. The results indicated that the accumulation of organic acid was a central adaptive mechanism by which L. quinquenervius maintained intracellular ionic balance under alkaline stress. Lathyrus quinquenervius may enhance organic acid synthesis to remedy the shortage of negative charge resulting from the massive influx of Na⁺ and decreased inorganic anions. In addition, saline stress and low alkaline stress slightly enhanced the activities of superoxide dismutase (SOD) and ascorbate peroxidase (APX), but did not affect catalase (CAT) activity. However, strong alkaline stress significantly enhanced the activities of SOD and APX, and reduced CAT activity. We propose that enhancing the activities of SOD and APX may be a vital mechanism by which L. quinquenervius resists oxidative stress caused by alkaline stress.     

Protective role of proline against salt stress is partially related to the improvement of water status and peroxidase enzyme activity in cucumber

A salt-sensitive cucumber cultivar "Jinchun No. 2" ( Cucumis sativus L.) was used to investigate the role of proline in alleviating salt stress in cucumber. Proline was applied twice (day 0 and day 4 after salt treatment) as a foliar spray, with a volume of 25 mL per plant at each time. Plant dry weight, leaf relative water content, proline, malondialdehyde (MDA), Na⁺, K⁺ and Cl⁻ contents, as well as superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) and ascorbate peroxidase (APX) activities in the plants were determined at day 8 after salt treatment. The results showed that 100 mmol L^–1 NaCl stress significantly decreased plant dry weight, leaf relative water and K⁺ contents, and increased leaf MDA, Na⁺ and Cl⁻ contents and SOD, POD, CAT and APX activities. However, leaf proline accumulation was not affected by salinity. The exogenous application of proline significantly alleviated the growth inhibition of plants induced by NaCl, and was accompanied by higher leaf relative water content and POD activity, higher proline and Cl⁻ contents, and lower MDA content and SOD activity. However, there was no significant difference in Na⁺ and K⁺ contents or in CAT and APX activities between proline-treated and untreated plants under salt stress. Taken together, these results suggested that the foliar application of proline was an effective way to improve the salt tolerance of cucumber. The enhanced salt tolerance could be partially attributed to the improved water status and peroxidase enzyme activity in the leaf.     

Interactive effect of salinity and silver nanoparticles on photosynthetic and biochemical parameters of wheat

The present study investigated the influence of seed priming with silver nanoparticles (Ag NPs), 0, 2, 5 and 10 mM, on growth and biochemical parameters of wheat (Triticum aestivum L.) under salt stress. As expected, 150 mM of NaCl decreased the shoot fresh and dry weights and chlorophyll contents and increased the catalase (CAT) and peroxidase (POD) activities. Salinity enhanced the concentration of proline, soluble sugars, malondialdehyde and hydrogen peroxide. Seed priming with Ag NPs increased the shoot fresh and dry weight of normal and salt-stressed plants. Lower concentration of Ag NPs decreased the total soluble sugars and proline contents, while the higher Ag NPs levels increased these contents compared to the control. The combined application of Ag NPs and salt stress increased the soluble sugars and proline contents, while it decreased CAT activity and increased POD activity compared to the respective Ag NPs treatments alone. Overall, our results demonstrated that Ag NPs enhanced the salt tolerance in wheat, but the long-term response of Ag NPs under salt stress needs further investigation.     

荆芥幼苗对盐胁迫的生理响应

为探究盐胁迫对荆芥幼苗生理特性的影响,采用盆栽砂培试验,研究不同浓度盐胁迫(0、25、50、75和100 mmol·L^-1 NaCl)下荆芥幼苗生长、质膜稳定性、渗透调节、抗氧化酶系统、离子吸收和分配的变化。结果表明,随着盐浓度的增加,荆芥盐害指数逐渐升高,幼苗株高增加速率和比叶面积均逐渐降低,单株干重和叶绿素含量均呈先增加后减少的趋势,且均在25 mmol·L^-1 NaCl处理下达到最大;叶片中丙二醛(MDA)含量和电解质渗漏率均显著增加;可溶性蛋白和脯氨酸含量均呈先增加后减少趋势,且分别在50和75 mmol·L^-1 NaCl处理下达到最大,而可溶性糖含量则不断上升;超氧化物歧化酶(SOD)、过氧化氢酶(CAT)活性均呈先上升后下降的趋势,在25 mmol·L^-1 NaCl处理下达到最大,而过氧化物酶(POD)活性则持续下降。盐胁迫下,荆芥的根、茎、叶大量积累Na^+,但主要集中在地上部分,同时各部位K^+、Ca²⁺含量及K^+/Na^+、Ca²⁺/Na^+值均显著下降。综上,荆芥幼苗对盐胁迫极为敏感,但对低浓度的盐胁迫(25 mmol·L^-1 NaCl)具有一定的耐受性。本研究结果为荆芥的规范化栽培和抗逆驯化研究奠定了理论基础。     

磷对盐碱胁迫下棉花离子平衡及相关调控基因的影响

[目的]盐胁迫是影响棉花生长的主要非生物胁迫之一,合理施肥促进盐胁迫下作物对养分离子的吸收,是提高作物耐盐性的重要途径.研究施磷随盐胁迫下棉花离子组的响应特征及Na+转运相关基因表达的变化,探讨磷对棉花耐盐性机制.[方法]采用盆栽试验,设置盐胁迫(NaCl)和碱胁迫(NaHCO3+Na2CO3)2个逆境处理,每个逆境下...     

外源24-表油菜素内酯对NaCl胁迫下桑树幼苗的缓解效应

为探究施加外源24-表油菜素内酯(EBR)对桑树耐盐性的影响,以桑品种桂桑优12为材料进行盆栽试验,在200 mmol·L^-1 NaCl胁迫下,研究外施不同浓度(0.01、0.10、1.00 μmol·L^-1)EBR对盐胁迫下桑树幼苗质量、根系特征、光合色素含量、活性氧(ROS)水平、丙二醛(MDA)含量、相对电导率、抗氧化酶活性和渗透调节物质含量等生理指标的影响。结果表明,与单一NaCl胁迫相比,外源添加0.10 μmol·L^-1EBR后,桑树幼苗地上鲜质量和干质量、主根长、总根长、根表面积、根平均直径、根总体积、叶绿素和类胡萝卜素含量均提高,叶片与根中产生羟基自由基(·OH)能力、产生超氧阴离子自由基($\text{O}_{2}^{\frac{{}}{\cdot }}$)活力、过氧化氢(H₂O₂)含量降低;叶片与根中的超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性升高;脯氨酸、可溶性蛋白和可溶性糖含量增加;相对电导率和MDA含量降低。其中地下部鲜质量和根体积是盐胁迫下的1.56和1.57倍,叶片中CAT活性和可溶性糖含量达到盐胁迫下的1.57和1.68倍。本研究为揭示外施油菜素内酯增强桑树耐盐性的生理调控机制提供了参考依据。     

番茄SlMIP基因参与转基因拟南芥的渗透调节

本实验以野生型（WT）和转番茄SlMIP基因的拟南芥为材料,研究NaCl胁迫条件下对二者体内渗透调节特征的影响。结果发现, 在NaCl胁迫下,转SlMIP基因的拟南芥细胞膜损伤程度较轻,组织渗透势显著降低,并保持较高的组织含水量,生长受抑制程度明显低于野生型植株。转基因植株体内Na+ 含量和Na+/K+ 比值显著低于野生型拟南芥,K+含量虽有所下降但仍显著高于野生型植株,而且在盐胁迫下脯氨酸和可溶性糖含量也高于野生型拟南芥,表明转入番茄SlMIP基因后的拟南芥,通过增加水分子的吸收,减少钠离子的进入,增加了细胞内脯氨酸和可溶性糖的含量,进而影响植物的有机渗透调节能力,与此同时可能通过离子化区隔机制以及与质膜Na+/H+逆向转运蛋白的相互作用更有效地调节细胞内外的无机离子交换能力,使植物更有效地抵御盐害,表明番茄SlMIP水通道蛋白基因在植物盐胁迫下具有重要的渗透调节作用。     

外源NO对盐胁迫下甜高粱种子萌发和幼苗生长的影响

为探究外源NO对盐胁迫下甜高粱种子萌发和幼苗生长的影响,以国能4号甜高粱为试验材料,采用不同浓度(0.05、0.1和0.2 mmol·L^-1)硝普钠(SNP,NO供体)处理1.6%NaCl胁迫下的种子和幼苗,统计种子发芽数,测定幼苗叶片叶绿素、脯氨酸、可溶性糖、可溶性蛋白、丙二醛(MDA)含量及超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)和抗坏血酸过氧化物酶(APX)活性等生理生化指标。结果表明,盐胁迫抑制了甜高粱种子的萌发,种子发芽指标(种子发芽率、发芽势和发芽指数)显著降低,1.6%NaCl是甜高粱种子萌发的敏感盐浓度;喷施不同浓度SNP可显著提高盐胁迫下甜高粱种子的发芽率及幼苗叶片叶绿素、可溶性糖、可溶性蛋白和脯氨酸含量,降低叶片MDA含量,增强抗氧化酶活性,其中0.1 mmol·L^-1SNP处理效果最佳。0.1 mmol·L^-1SNP处理下,与单独盐浓度处理相比,甜高粱种子发芽率、发芽势和发芽指数分别增加了29.51%、39.21%和38.91%;幼苗叶片叶绿素a、叶绿素b、总叶绿素、可溶性糖、可溶性蛋白和脯氨酸含量分别提高了230.00%、184.38%、214.13%、17.00%、8.78%和40.63%;MDA含量降低了34.01%;SOD、POD、CAT和APX活性分别增强了33.38%、55.75%、23.17%和116.46%。综上,盐胁迫下适宜浓度的SNP处理,可提高甜高粱幼苗叶片渗透调节物质含量,增强抗氧化酶活性,清除体内活性氧,从而促进幼苗生长,增强植株抗盐性。本研究结果为提高甜高粱耐盐性及揭示其耐盐机制提供了理论依据。     

Interaction forces between soil particles: shear moduli of the < 2 μm size fraction

The shear moduli of the < 2 μm size fraction of three soils have been measured for samples of 10–35% w/w solid. Samples were thixotropic, the shear modulus increasing with time. This increase can be described by simple models and visualized in terms of the formation of links between particles. For a given soil the shear modulus increases with ionic form in the order Ca²⁺∼Mg²⁺ + + < Li⁺ and varies in a complex manner with electrolyte concentration.     

Ca2+ Effects on K+ Fluxes in Arabidopsis Seedlings Exposed to Al3+

Potassium transport was investigated in the root elongation zone of Arabidopsis seedlings during the first minutes of Al³⁺ exposure, using the non-invasive MIFE microelectrode technique. To prevent pH changes during Al³⁺ application, and to separate aluminium from acidic stress, plants were pre-treated with 5 mM homoPIPES before addition of AlCl₃ (pH 4.2). The 30-min treatment with 50 or 500 μM AlCl₃ led to a significant increase in K⁺ efflux in solutions containing 100 μM CaCl₂. This efflux was suppressed by high concentrations of Ca²⁺ (10 mM) in the bathing solution. Our results suggest that elevated external Ca²⁺ activities can sustain K⁺ influx in the root elongation zone during Al³⁺ exposure either by maintaining [Ca²⁺]_cyt or by affecting Al³⁺ uptake across the plasma membrane.     

Physiological Responses of Leymus Chinensis to Long-Term Salt,Alkali and Mixed Salt-Alkali Stresses

The aim of this study was to evaluate the physiological responses of Leymus chinensis (Trin.) Tzvel exposed to long-term salt, salt-alkali, and alkali stress in order to elucidate how L. chinensis can survive under alkaline-sodic soils. L. chinensis (30 days after germination) were stressed with salt [SS; sodium chloride (NaCl)], mixed salt-alkali [MS; molar ratio of NaCl: sodium carbonate (Na₂CO₃) = 2:1] and alkali salt (AS; Na₂CO₃) at four different levels of sodium (Na⁺) concentration (0, 75, 150, and 300 mM) for 60 days. L. chinensis showed 100% survival rate at all treatments except 300 mM SS (33.3%) and AS (18.9%). The growth and physiological parameters of survival plants were measured. As anticipated, growth of L. chinensis was inhibited after stresses, which reflected in the decline of plant height, dry weight and tiller number following the increased Na⁺ concentration. The content of Na⁺, proline, and soluble sugar in L. chinensis increased with the increasing Na⁺ concentration, suggesting that L. chinensis need to accumulate inorganic and organic solutes for resisting osmotic stress induced by various salt stresses. These processes ensure the water balance that can provide a relative normal physiological environment for L. chinensis. Potassium (K⁺) content of L. chinensis kept at a relative lower level than control to ensure the normal physiological processes. Chlorophyll content of stressed plant increased slightly compared to control plants, which can produce more energy for L. chinensis resisting various stresses. The increased malondialdehyde (MDA) content of stressed plants showed the damage of various stresses. Among the three treatments (SS, MS, and AS), the injury extent for L. chinensis can be expressed by AS>SS>MS, and MS was the most complicated for the counterbalance effects of soil electrical conduction and pH.     

羊草对盐碱胁迫的生理生化响应特征

研究羊草对盐碱胁迫生理响应特征,旨在找出羊草耐盐碱范围及其对盐碱地改良的应用潜力。通过模拟5组不同盐碱梯度土壤胁迫试验(p H值为7.61、8.05、8.33、8.73、9.37,电导率为0.109、0.301、0.437、0.486、0.591 m S/cm),研究了羊草体内离子含量和运输能力、有机小分子渗透调节和保护酶活性变化。结果表明,盐碱胁迫由低到高,超氧化物歧化酶和甜菜碱在中度盐碱胁迫(p H值8.33,电导率0.437 m S/cm)积累最多,其他测定的耐盐碱物质,钠离子、脯氨酸、丙二醛、有机酸(草酸、柠檬酸、酒石酸)、过氧化物酶、过氧化氢酶、可溶性总糖、可溶性蛋白均呈直线增加。在盐碱胁迫条件下,羊草叶片中积累的钾离子、甜菜碱、有机酸、可溶性蛋白高于根茎,而脯氨酸、丙二醛、保护酶(超氧化物歧化酶、过氧化物酶、过氧化氢酶)、可溶性总糖叶片中含量低于根茎。根茎大量积累保护酶、脯氨酸及丙二醛有助于防止盐碱环境下根茎中膜过氧化,并保持根茎正常的渗透调节能力,从而有利于将K+等养分运送到叶片。在盐碱胁迫下,羊草叶片主要积累甜菜碱和有机酸进行渗透调节。该试验结果表明,羊草根茎和叶片在不同盐碱胁迫条件下各耐盐碱生理指标均产生一系列适应性反应,使其在盐碱环境正常生长,进而降低土壤盐碱性。该研究可为耐盐碱饲草选育及根茎型禾草改良盐碱化草地提供理论和实践依据。     

Exchangeable potassium and potassium balances in organic crop rotations on a coarse sand

Abstract. Crops on sandy soils (<5% clay) are exposed to K deficiency due to the small release and high leaching losses of K. Reliable tools are needed to improve the K management in cropping systems with limited K input, such as organic farming where import of nutrients are restricted according to the EC regulations. We investigated K balances and exchangeable K (K_exch) changes in an organic crop rotation experiment. Potassium leaching decreased from 42 kg ha⁻¹ in 1998/99 to 21 kg ha⁻¹ in 2000/01 as an average of a crop rotation (spring barley, grass-clover, winter wheat and pea/barley) with manure application and without catch crops. In the same period, spring K_exch decreased from 5.0 to 3.0 mg K 100 g soil⁻¹ (0–20 cm). The retention of the straw K left in the field after harvest increased with decreasing levels of K_exch. The cereal crops did not respond to K application but in the pea/barley mixture the pea yield increased by 46%. The concordance between measured K balances and changes in K_exch was weak. Exchangeable K is suitable as a tool for K management on a rotational basis, and a K_exch above 3 mg 100 g soil⁻¹ in the autumn should be avoided to minimize K leaching.     

种子引发对NaCl胁迫下小麦幼苗生理特性的影响

以耐盐性强的晋农207和耐盐性弱的运麦218两个冬小麦品种种子为试材,用20%的PEG及100mmol/L的KNO3水溶液对种子进行引发处理.研究模拟NaCl胁迫环境下引发处理对小麦种子发芽、幼苗耐盐性及幼苗生理特性的影响.结果表明:两种引发处理不同程度地提高了2个品种小麦的发芽势、发芽率、发芽指数和活力指数;使2个小...     

Influence of arbuscular mycorrhizae on yield,nutrients, organic solutes,and antioxidant enzymes of two wheat cultivars under salt stress

The efficacy of arbuscular mycorrhizae (AM) on nutrients, organic solutes, and antioxidant enzymes of wheat under salt stress was investigated and related to root colonization and plant productivity. The mycorrhizal inoculation increased N, P, K, Ca, and Mg uptake, soluble sugars, free amino acids, and proline accumulation, as well as peroxidase and catalase activities under saline conditions as compared to nonmycorrhizal plants. On the other hand, Na concentration was lower in mycorrhizal than in nonmycorrhizal plants grown under saline conditions. Arbuscular mycorrhizae protected wheat against the detrimental effects of salinity and stimulated its productivity. Hence, mycorrhizal colonization can play a vital role in the mitigation of the adverse effects of salinity by improving the wheat osmotic adjustment response, enhancing its defense system, and alleviating oxidative damage to cells. Arbuscular mycorrhizae are able to alter plant physiology in a way that empowers the plant to grow more efficiently on salt‐affected lands.     

Is burial of wheat straw in ditches a way to reduce CH4 emissions from rice cultivation?

Burial of wheat straw in ditches and incorporation of wheat straw are the two main ways of returning wheat straw prior to rice cultivation in China. To examine the effect of burying wheat straw in ditches on CH₄ emissions from rice cultivation, a field experiment was conducted at Yixing, Jiangsu, China in 2004. CH₄ flux was measured using a closed-chamber technique in three treatments (CK, no wheat straw application; WI, evenly incorporating 3.75 t ha⁻¹ wheat straw into the 0.1 m topsoil; WD, burying 3.75 t ha⁻¹ wheat straw in 0.14-m deep by 0.25-m wide ditches). Seasonal CH₄ emissions ranged from 49.7 to 218.4 kg CH₄ ha⁻¹. The application of wheat straw in these two ways significantly increased CH₄ emissions by 4.0-fold and 4.4-fold, respectively ( P  < 0.05). Although CH₄ flux from the non-ditch area in the WD treatment was as low as that in the CK treatment, it was counter-balanced by extremely high CH₄ flux from the ditch, which was approximately 6.0-fold as much as that from WI, leading to comparability between treatments WI and WD in total CH₄ emissions ( P  > 0.05). No significant difference was observed between the three treatments in grain yield ( P  > 0.05). The results indicated that burial of wheat straw in ditches is not a way to reduce CH₄ emission from rice cultivation.     

水分胁迫下脱落酸及磷脂酶在玉米幼苗根系渗透调节物质积累中的信号作用

试验研究了水分胁迫下、脱落酸 (ABA)、PLD/PLC抑制剂新霉素硫酸盐及脱落酸合成抑制剂Ancymidol在玉米幼苗根系渗透调节物质积累中的信号作用 ,结果表明水分胁迫下脱落酸可促进脯氨酸、可溶性糖和Ca2 的积累 ,但不能促进游离氨基酸和K 的积累 ,且PLD/PLC可能对脯氨酸、可溶性糖和游离氨基酸的积累有一定促进作用 ,但对K 和Ca2 的积累不起作用     

Correlations between extractable Na, K, Mg, Ca, P & N from fresh and dried samples of two Aquults

Significant increases in extractable ions resulted from air-drying and grinding samples of two infertile Aquults. Effects of the sample preparation differed markedly between ions and between the two soils. Regression equations were calculated to predict extractable ions in dried, ground samples from extractable ions in fresh, unground samples and the relationships were compared between the two soil series. Regressions were significantly different between soils for extractable PO³₄, Mg⁺⁺, and K⁺, but not for Ca⁺⁺ and Na⁺. Extractable NH ⁺₄ and NO-₃ in fresh, unground samples were not correlated with those in air-dry, ground samples of either soil. Differences in response to preparation between soil types appeared to be related to the oxidative status of these soils in the field, wherein constituents of more poorly-drained soils may be less stable to the oxidizing conditions of air-drying and grinding. Such complexities suggest that effects of sample preparation should be considered when interpreting soil nutrient data for studies of forest nutrient cycling and forest soil fertility.     

Accumulation,distribution, and physiological contribution of oxalic acid and other solutes in an alkali‐resistant forage plant,Kochia sieversiana,during adaptation to saline and alkaline conditions

Kochia sieversiana (Pall.) C. A. Mey is a forage plant in the family Chenopodiaceae, which can grow in extremely alkalinized grasslands at pH levels of 10 or higher. Kochia sieversiana often contains a large amount of oxalic acid. In the present study, seedlings of K. sieversiana were exposed to the following conditions: nonstress, salt stress (molar ratio of NaCl : Na₂SO₄ = 1:1, salinity: 200 mM), and alkali stress (molar ratio of NaHCO₃ : Na₂CO₃ = 1:1, salinity: 200 mM). By determining and analyzing various physiological factors such as the concentrations and distribution of different organic acids (including oxalic acid) in various parts of K. sieversiana, the concentrations of inorganic ions (K⁺, Na⁺, Cl^–, SO$ _4^{2-} $ , etc.), the organic solutes (proline, betaine, and soluble sugar) in shoots, and the accumulation and distribution of oxalic acid in K. sieversiana, the physiological contribution of oxalic acid to K. sieversiana adaptability to saline and alkaline conditions was investigated. Results show that oxalic acid mainly accumulated in shoots, and that its concentration was highest in mature functional leaves where photosynthesis productivity was based and lowest in old stems and roots, regardless of plant treatment (nonstress, salt, or alkali conditions). Under nonstress, salt, and alkali conditions, the concentrations of oxalic acid in mature leaves were 8%, 10%, and 12% of their dry weights, respectively, and were 1%, 0.7%, and 0.6% of dry weights, respectively, in roots. There were varying effects of salt and alkali conditions on oxalic acid concentrations in different parts of K. sieversiana. Oxalic acid concentration increased in leaves, did not change significantly in young stems, and decreased in old stems and roots. The present analysis shows that oxalic acid exists as an organic anion in K. sieversiana. Consequently, oxalic acid not only plays a crucial role in osmoregulation and pH adjustment, but it also is the dominant contributor of negative charge, playing a key role in maintaining ionic balance in vivo. Oxalic acid in K. sieversiana shoots is a key substance on which the adaptation to saline and alkaline conditions is based.