Production of Polyamines Is Enhanced by Endogenous Abscisic Acid in Maize Seedlings Subjected to Salt Stress

It is known that salt stress and exogenously applied abscisic acid （ABA） can enhance the polyamine content in plants and that salt stress itself can lead to an increase in endogenous ABA production. In the present study, the relationships between salt-induced ABA and polyamine accumulation were inves- tigated using ABA-deficient mutant （vp5/vp5） maize （Zea mays L.） seedlings and ABA and polyamine biosynthesis inhibitors. The results show that reduced endogenous ABA levels, as a result of either the mutation or by using a chemical inhibitor （sodium tungstate）, also reduced the accumulation of polyamines in salt-stressed leaves of maize seedlings. The polyamine synthesis inhibitors D-arginine and α- difluoromethylornithine also reduced the polyamine content of the leaves of maize seedling under salt stress. Both ABA and polyamine enhanced the dry weight accumulation of salt-stressed seedlings and also increased the activities of the two dominant tonoplast membrane enzymes, H^＋-ATPase and H^＋-PPase, when plants were under salt stress. The results suggest that salt stress induces an increase in endogenous ABA levels, which then enhances polyamine synthesis. Such responses may increase a plant＇s tolerance to salt.     

Ectopic Expression of a Bacterium NhaD-type Na~+/H~+ Antiporter Leads to Increased Tolerance to Combined Salt/Alkali Stresses

AaNhaD,a gene isolated from the soda lake alkaliphile Alkalimonas amylolytica,encodes a Na+/H+ antiporter crucial for the bacterium’s resistance to salt/alkali stresses.However,it remains unknown whether this type of bacterial gene may be able to increase the tolerance of flowering plants to salt/alkali stresses.To investigate the use of extremophile genetic resources in higher plants,transgenic tobacco BY-2 cells and plants harboring AaNhaD were generated and their stress tolerance was evaluated.Ectopic expression of AaNhaD enhanced the salt tolerance of the transgenic BY-2 cells in a pH-dependent manner.Compared to wild-type controls,the transgenic cells exhibited increased Na+concentrations and pH levels in the vacuoles.Subcellular localization analysis indicated that AaNhaD-GFP fusion proteins were primarily localized in the tonoplasts.Similar to the transgenic BY-2 cells,AaNhaD-overexpressing tobacco plants displayed enhanced stress tolerance when grown in saline-alkali soil.These results indicate that AaNhaD functions as a pH-dependent tonoplast Na+/H+antiporter in plant cells,thus presenting a new avenue for the genetic improvement of salinity/alkalinity tolerance.     

Seed Treatment with Auxins Modulates Growth and Ion Partitioning in Salt-stressed Wheat Plants

Experiments were performed to determine whether seed priming with different concentrations （100, 150, and 200 mg/L） of auxins （indoleacetic acid （IAA）, indolebutyric acid （IBA）, or their precursor tryptophane （Trp）） could alter salinity induced perturbances in salicylic acid and ion concentrations and, hence, growth in wheat （Triticum aestivum L.） cultivars, namely M.H.-97 （salt intolerant） and tnqtab-91 （salt tolerant）. Primed and non-primed seeds were sown in Petri dishes in a growth room, as well as in a field treated with 15 dS/m NaCl salinity. All priming agents, except IBA, increased the final germination percentage in both cultivars. The seedlings of either cultivar raised from Trp-treated seeds had greater dry biomass when under salt stress. In field experiments, Trp priming was much more effective in mediating the increase in grain yield, irrespective of the cultivar, under salt stress. The alleviatory effect of Trp was found to be associated with reduced uptake of Na^＋ in the roots and subsequent translocation to the shoots, as well as increased partitioning of Ca^＋ in the roots of salt-stressed wheat plants. Plants of both cultivars raised from Trp-and IAA-treated seeds accumulated free salicylic acid in their leaves when under salt stress. Overall, the Trp priming-induced improvement in germination and the subsequent growth of wheat plants could be related to ion homeostasis when under salt stress. The possible involvement of salicylic acid in the Trp priming-induced better growth under Conditions of salt stress is discussed.     

Hydrogen Peroxide in Plants： a Versatile Molecule of the Reactive Oxygen Species Network

Plants often face the challenge of severe environmental conditions, which include various biotic and abiotic stresses that exert adverse effects on plant growth and development. During evolution, plants have evolved complex regulatory mechanisms to adapt to various environmental stressors. One of the consequences of stress is an increase in the cellular concentration of reactive oxygen species （ROS）, which are subsequently converted to hydrogen peroxide （H2O2）. Even under normal conditions, higher plants produce ROS during metabolic processes. Excess concentrations of ROS result in oxidative damage to or the apoptotic death of cells. Development of an antioxidant defense system in plants protects them against oxidative stress damage. These ROS and, more particularly, H2O2, play versatile roles in normal plant physiological processes and in resistance to stresses. Recently, H2O2 has been regarded as a signaling molecule and regulator of the expression of some genes in cells. This review describes various aspects of H2O2 function, generation and scavenging, gene regulation and cross-links with other physiological molecules during plant growth, development and resistance responses.     

Improvement of plant abiotic stress tolerance through modulation of the polyamine pathway FA简

Polyamines （mainly putrescine （Put）, spermidine （Spd）, and spermine （Spin）） have been widely found in a range of physiological processes and in almost all diverse environ- mental stresses. In various plant species, abiotic stresses modulated the accumulation of polyamines and related gene expression. Studies using loss-of-function mutants and transgenic overexpression plants modulating polyamine metabolic pathways confirmed protective roles of polyamines during plant abiotic stress responses, and indicated the possibility to improve plant tolerance through genetic manipulation of the polyamine pathway. Additionally, puta- tive mechanisms of polyamines involved in plant abiotic stress tolerance were thoroughly discussed and crosstalks among polyamine, abscisic acid, and nitric oxide in plant responses to abiotic stress were emphasized. Special attention was paid to the interaction between polyamine and reactive oxygen species, ion channels, amino acid and carbon metabolism, and other adaptive responses. Further studies are needed to elucidate the polyamine signaling pathway, especially polyamine-regulated downstream tar- gets and the connections between polyamines and other stress responsive molecules.     

Seed Treatment with Auxins Modulates Growth and Ion Partitioning in Salt-stressed Wheat Plants

Experiments were performed to determine whether seed priming with different concentrations (100, 150, and 200 mg/L) ofauxins (indoleacetic acid (IAA), indolebutyric acid (IBA), or their precursor tryptophane (Trp)) could alter salinity inducedperturbances in salicylic acid and ion concentrations and, hence, growth in wheat (Triticum aestivum L.) cultivars, namelyM. H.-97 (salt intolerant) and Inqlab-91 (salt tolerant). Primed and non-primed seeds were sown in Petri dishes in a growthroom, as well as in a field treated with 15 dS/m NaCl salinity. All priming agents, except IBA, increased the final germinationpercentage in both cultivars. The seedlings of either cultivar raised from Trp-treated seeds had greater dry biomass whenunder salt stress. In field experiments, Trp priming was much more effective in mediating the increase in grain yield,irrespective of the cultivar, under salt stress. The alleviatory effect of Trp was found to be associated with reduced uptakeof Na~ in the roots and subsequent translocation to the shoots, as well as increased partitioning of Ca~(2 )in the roots ofsalt-stressed wheat plants. Plants of both cultivars raised from Trp-and IAA-treated seeds accumulated free salicylic acidin their leaves when under salt stress. Overall, the Trp priming-induced improvement in germination and the subsequentgrowth of wheat plants could be related to ion homeostasis when under salt stress. The possible involvement of salicylicacid in the Trp priming-induced better growth under conditions of salt stress is discussed.     

Physiological Responses of Zea mays Seedlings to Interactions Between Cadmium and Salinity

The effects of 0, 2.5, 5.0, and 10.0 pmol/L Cd^2＋（Cd（NO3）2.4H2O） and 0, 10, 25, 50, and 100 mmol/L NaCl on growth, photosynthesis and the content of some ions in maize （Zea mays L.） were investigated in the present study. With Increasing concentrations of Cd^2＋ or NaCI alone in Hoagland nutrient solution, the chlorophylls and starch content decreased. Combination treatment with salinity and cadmium increased the negative effects observed following the two stresses alone. Plants exhibiting growth retardation in response to one mild stress factor （25-50 mmol/L NaCl） became more tolerant to the other stress factor （Cd）. The exposure of plants to cadmium caused a partial reversal of the effects of salinity. Root and shoot growth, ion accumulation and levels of photosynthetic pigments were improved at moderate concentrations of the two stress factors Imposed jointly.     

Na~ and Water Uptake in Relation to the Radial Reflection Coefficient of Root in Arrowleaf Saltbush Under Salt Stress

The response of halophyte arrowleaf saltbush(Atriplex triangularis Willd)plants to a gradient of salt stress were investigatedwith hydroponically cultured seedlings.Under salt stress,both the Na~ uptake into root xylem and negative pressures inxylem vessels increased with the elevation of salinity(up to 500 mol/m~3)in the root environment.However,the increment innegative pressures in root xylem far from matches the decrease in the osmotic potential of the root bathing solutions,evenwhen the osmotic potential of xylem sap is taken into consideration.The total water potential of xylem sap in arrowleafsaltbush roots was close to the osmotic potential of root bathing solutions when the salt stress was low,but a progressivelyincreased gap between the water potential of xylem sap and the osmotic potential of root bathing solutions was observedwhen the salinity in the root environment was enhanced.The maximum gap was 1.4 MPa at a salinity level of 500 mol/m~3without apparent dehydration of the tested plants.This discrepancy could not be explained with the current theories inplant physiology.The radial reflection coefficient of root in arrowleaf saltbush decreased with the enhanced salt stress wasand accompanied by an increase in the Na~ uptake into xylem sap.However,the relative Na~ in xylem exudates based onthe corresponding NaCl concentration in the root bathing solutions showed a tendency of decrease.The results showedthat the reduction in the radial reflection coefficient of roots in the arrowleaf saltbush did not lead to a mass influx of NaClinto xylem when the radial reflection coefficient of the root was considerably small;and that arrowleaf saltbush could usesmall xylem pressures to counterbalance the salt stresses,either with the uptake of large amounts of salt,or with thedevelopment of xylem pressures dangerously negative.This strategy could be one of the mechanisms behind the highresistance of arrowleaf saltbush plants to salt stress.     

Does Seed Priming Induce Changes in the Levels of Some Endogenous Plant Hormones in Hexaploid Wheat Plants Under Salt Stress？

In order to assess whether salt tolerance could be Improved In spring wheat （Triticum aestivum L.）, the present study was performed by soaking the seeds of two cultlvars, namely MH-97 （salt sensitive） and Inqlab-91 （salt tolerant）, for 12 h In distilled water or 100 mol/m^3 CaCl2, KCI, or NaCI. Primed seeds from each treatment group and non-primed seeds were sown In a field In which NaCI salinity of 15 dS/m was developed. Priming of seeds with CaCl2, followed by priming with KCI and NaCI, was found to be effective In alleviating the adverse effects of salt stress on both wheat cultivars In terms of shoot fresh and dry weights and grain yield. Priming with CaCl2 alleviated the adverse effects of salt stress on hormonal balance In plants of both cultlvars. In MH-97 plants, CaCl2 pretreatment considerably reduced leaf absclslc acid （ABA） concentrations and Increased leaf free salicylic acid （SA） concentrations under both saline and non-saline conditions. In contrast, In the Inqlab-91 plant, CaCl2 Increased free Indoleacetic acid （IAA） and indolebutyrlc acid （IBA） content. However, priming of seeds with CaCl2 did not alter free polyamlne levels in either cultlvar, although spermldlne levels were considerably lower In plants raised from seeds treated with CaCl2 for both cultlvars under saline conditions. Priming with KCI Increased growth In Inqlab-91 plants, but not In MH-97 plants, under saline conditions. The salinity Induced reducUon In auxins （IAA and IBA） was alleviated by NaCI priming In both cultlvars under saline conditions. However, NaCI Increased leaf free ABA content and lowered leaf SA and putresclne levels In Inqlab-91 plants under saline conditions. In conclusion, although all three priming agents （I.e. CaCl2, KCI, and NaCI） were effective In alleviating the adverse effects of salt stress on wheat plants, their effects on altering the levels of different plant hormones were different In the two cuItlvars.     

Effects of various mixed salt-alkaline stresses on growth,photosynthesis, and photosynthetic pigment concentrations of <Emphasis Type="Italic">Medicago ruthenica</Emphasis> seedlings

Soil salinization and alkalinization frequently co-occur in naturally saline and alkaline soils. To understand the characteristics of mixed salt-alkali stress and adaptive response of Medicago ruthenica seedlings to salt-alkali stress, water content of shoots, growth and photosynthetic characteristics of seedlings under 30 salt-alkaline combinations (salinity 24–120 mM and pH 7.03–10.32) with mixed salts (NaCl, Na₂SO₄, NaHCO₃, and Na₂CO₃) were examined. The indices were significantly affected by both salinity and pH. The interactive effects between salt and alkali stresses were significant, except for photosynthetic pigments. Water content of shoots, relative growth rates of shoots and roots and pigment concentrations showed decreasing trends with increasing salinity and alkalinity. The root activity under high alkalinity and salinity treatments gradually decreased, but was stimulated by the combined effects of low alkalinity and salinity. The survival rate decreased with increased salinity, except at pH 7.03–7.26 when all plants survived. Net photosynthetic rate, stomatal conductance and intercellular CO₂ concentration decreased with increased salinity and pH. M. ruthenica tolerated the stress of high salt concentration when alkali concentration was low, and the synergistic effects of high alkali and high salt concentrations lead to the death of some or all seedlings. M. ruthenica appeared to be saltalkali tolerant. Reducing the salt concentration or pH based on the salt components in the soil may be helpful to abate damage from mixed salt-alkaline stress.     

盐碱协同胁迫对向日葵抗氧化酶系统的影响

根据中国东北盐碱土壤特点,将4种盐NaCl、NaHCO3、Na2SO4和Na2CO3按不同比例混合,模拟出25种盐度和pH值各不相同的复杂盐碱条件(盐浓度为50～250 mmol/L,pH值为712～1046),并对向日葵苗进行盐碱混合胁迫处理,研究了向日葵超氧化物歧化酶(SOD)、过氧化氢酶(CAT)和过氧化物酶(POD)酶等抗氧化酶系统和丙二醛(MDA)的盐碱协同胁迫效应．结果表明, 向日葵抗氧化物酶活性强弱同时与盐度和碱度密切相关,3种抗氧化物酶活性对于盐浓度的反应相似,均为其含量随着盐浓度的升高开始逐渐升高然后下降,而对于pH的影响,不同酶反应结果不同．即随着pH值升高,SOD酶活性和CAT酶活性降低,而POD酶活性反应则是随着pH值升高活性也升高．双向方差分析(ANOVA)结果表明：盐碱效应对于3种酶活力的影响是显著的．其中,盐效应对POD和SOD活性的影响比pH值的影响大,而pH值对CAT活性的影响效应比盐效应大．除SOD外,盐碱效应的交互作用显著 (P<0001)．抗氧化酶系统和MDA含量两者间相关性和逐步回归分析表明,3种酶对MDA的影响效应随其强度不同呈现显著不同．其中SOD是1个主导因子,CAT 处于次位, 而POD的影响不大,甚至可以忽略．     

Effects of various salt-alkaline mixed stresses on Aneurolepidium chinense (Trin.) Kitag.

The stress conditions of salt-alkalinized soil were simulated to investigate the features and acting factors of salt-alkaline mixed stress, using a natural salt-alkaline tolerant grass Aneurolepidium chinense (Trin.) Kitag. According to the features of salt-alkalinized soil in the northeast of China, various salt-alkali conditions with different salinities and pHs were established by mixing NaCl, NaHCO₃, Na₂SO₄, and Na₂CO₃, in various proportions. The treatments included a salt concentration range of 50 to 350 mM and pH values from 7.14 to 10.81. Seedlings of A. chinense were stressed under these salt-alkali conditions. Several physiological indices of seedling stress were determined, including survival rate, tillering rate, number of rhizomes, relative growth rate (RGR), proline content, electrolyte leakage rate, and Na⁺ and K⁺ content, in order to analyze the characteristics of the stresses due to the salt-alkali mixes and their main stress factors.The results showed that the survival rate, tillering rate, number of rhizomes, RGR, and K⁺ content of A. chinense decreased with increasing salinity and pH (or alkalinity). Proline and Na⁺ content and electrolyte leakage rate increased with increasing salinity and pH (or alkalinity). The deleterious effects of a high pH value or salinity alone were significantly less than those of high pH in combination with salinity. This result suggested that for a salt-alkaline mixed stress, a reciprocal enhancement between salt stress and alkali stress was a characteristic feature, and it was most evidently reflected in the survival rate. When salinity was below 125 mM or pH was below 8.8, survival rates were all 100%. However, when salinity was above 125 mM and pH was above 8.8, survival rates sharply declined with the increasing of either salinity or pH.The buffer capacity of the treatment solution was taken as a stress factor in order to simplify the stress factor analysis. The results of the statistical analysis showed that for the stress factors of the salt-alkaline mixed stress, [CO₃^2–] and [HCO₃^–] could be fully represented by the buffer capacity, and [Na⁺] could be fully represented by salinity, whereas [SO₄^2–] was negligible. Therefore, four factors, salinity, buffer capacity, pH and [Cl^–], could reflect all of the stress factors. Perfect linear correlations were observed between all physiological indices and four or three stress factors by a stepwise regression analysis. However, the effects of the four stress factors on the physiological indices were significantly different in magnitude. Buffer capacity and salinity were dominant factors for all physiological indices. Thus, it is reasonable to consider the sum of salinity plus buffer capacity as the strength value of salt-alkaline mixed stress. Furthermore, the relationships between different physiological indices and various stress factors were shown to be different.     

NaCl和Na2CO3胁迫对桑树幼苗生长和光合特性的影响

以1年生“青龙桑”幼苗为试验材料,研究了中性盐(NaCl)和碱性盐(Na2CO3)胁迫下桑树幼苗的生长和叶片光合特性.结果表明:盐胁迫明显降低了桑树幼苗的株高、叶片数、生物量和叶片的光合能力.随着Na+浓度的增加,桑树叶片的气孔导度、蒸腾速率、净光合速率、实际光化学效率、电子传递速率和光化学猝灭系数明显降低,过剩光能以非光化学猝灭形式耗散的比例增加,桑树叶片的光能转化效率和光合能力下降.在Na+浓度＜150 mmol·L-1时,桑树幼苗的光合能力和生长受到的抑制较小,通过增加根冠比进一步适应盐胁迫,但这种保护机制随着盐浓度的增加逐渐降低.在Na2CO3胁迫下,＞50 mmol·L-1 Na+浓度对桑树的生长和光合能力表现出较强的抑制作用,并随Na+浓度的增加,抑制程度加大.在NaCl＜ 150mmol·L-1时,桑树的光合能力主要依赖植株形态和光合代谢双重途径适应中性盐逆境,而在NaC1浓度＞150 mmol·L-1和碱性盐胁迫下,其主要依赖光合代谢来适应逆境.     

NaCl和Na2CO3胁迫对桑树幼苗生长和光合特性的影响

以1年生“青龙桑”幼苗为试验材料,研究了中性盐(NaCl)和碱性盐(Na2CO3)胁迫下桑树幼苗的生长和叶片光合特性.结果表明： 盐胁迫明显降低了桑树幼苗的株高、叶片数、生物量和叶片的光合能力.随着Na+浓度的增加,桑树叶片的气孔导度、蒸腾速率、净光合速率、实际光化学效率、电子传递速率和光化学猝灭系数明显降低,过剩光能以非光化学猝灭形式耗散的比例增加,桑树叶片的光能转化效率和光合能力下降.在Na+浓度＜150 mmol·L-1时,桑树幼苗的光合能力和生长受到的抑制较小,通过增加根冠比进一步适应盐胁迫,但这种保护机制随着盐浓度的增加逐渐降低.在Na2CO3胁迫下,＞50 mmol·L-1Na+浓度对桑树的生长和光合能力表现出较强的抑制作用,并随Na+浓度的增加,抑制程度加大.在NaCl＜150 mmol·L-1时,桑树的光合能力主要依赖植株形态和光合代谢双重途径适应中性盐逆境,而在NaCl浓度＞150 mmol·L-1和碱性盐胁迫下,其主要依赖光合代谢来适应逆境.     

Photosynthesis,chlorophyll fluorescence,inorganic ion and organic acid accumulations of sunflower in responses to salt and salt-alkaline mixed stress

Sunflowers were treated with mixing proportions of NaCl, Na₂SO₄, NaHCO₃, and Na₂CO₃. Effects of salt and saltalkaline mixed stress on growth, photosynthesis, chlorophyll fluorescence, and contents of inorganic ions and organic acids of sunflower were compared. The growth of sunflower decreased with increasing salinity. The contents of photosynthetic pigments did not decrease under salt stress, but their contents decreased sharply under salt-alkaline mixed stress. Net photosynthetic rates, stomatal conductance and intercellular CO₂ concentration decreased obviously, with greater reductions under salt-alkaline mixed stress than under salt one. Fluorescence parameters showed no significant differences under salt stress. However, maximal efficiency of PSII photochemistry, photochemical quenching coefficient, electron transport rate, and actual PSII efficiency significantly decreased but non-photochemical quenching increased substantially under salt-alkaline mixed stress. Under salt-alkaline mixed stress, sunflower leaves maintained a low Na⁺- and high K⁺ status; this may be an important feature of sunflower tolerance to salinity. Analysis of the mechanism of ion balance showed that K⁺ but not Na⁺ was the main inorganic cation in sunflower leaves. Our results indicated that the change in organic acid content was opposite to the change of Cl⁻, and the contribution of organic acid to total charge in sunflower leaves under both stresses decreased with increasing salinity. This may be a special adaptive response to stresses for sunflower. Sunflower under stress conditions mainly accumulated inorganic ions instead of synthesizing organic compounds to decrease cell water potential in order to save energy consumption.     

Interactive effects of salt and alkali stresses on seed germination,germination recovery,and seedling growth of a halophyte Spartina alterniflora (Poaceae)

Soil salinization and alkalinization frequently co-occur in nature, but there is little information on the interactive effects of salt and alkali stresses on plants. Seed germination and early seedling growth are crucial stages for plant establishment. We investigated the interactive effects of salt and alkali stresses on seed germination, germination recovery and seedling growth of a halophyte Spartina alterniflora. Seed germination percentage was not significantly reduced at low salinity (≤ 200 mM) at pH 6.63–9.95, but decreased with increased salinity and pH. Ungerminated seeds germinated well after transfer to distilled water from treatment solutions, indicating that seeds can remain viable in high salt–alkaline habits. Shoot growth was stimulated at low salinity and pH, but decreased with increased salinity and pH. Radicle elongation decreased sharply with increased salinity and pH, and was significantly inhibited when pH ≥ 9.0, indicating that the radicles are very sensitive to salt–alkaline stress. The deleterious effects of salinity or high pH alone were less than when combined. A reciprocal enhancement of salt and alkali stresses is a characteristic feature for salt–alkaline stress. Stepwise regression analysis indicates that salinity is the dominant factor, while pH and buffer capacity are secondary for salt–alkaline mixed stress.     

盐、碱胁迫下小冰麦体内的pH及离子平衡

通过混合两种中性盐（NaCl和Na₂SO₄）和两种碱性盐（NaHCO₃和Na₂CO₃）分别模拟出不同强度的盐、碱胁迫条件，对小冰麦苗进行12 d胁迫处理，测定茎叶组织液的pH值及Na⁺、K⁺、Ca²⁺、Cl^-、SO₄^2-、NO₃^-、H₂PO₄-和有机酸等溶质的浓度，以探讨盐、碱两种胁迫下小冰麦体内的pH及离子平衡特点.结果表明：盐、碱胁迫下小冰麦茎叶内的pH值均稳定不变；随胁迫强度的增加，盐胁迫下小冰麦茎叶内有机酸浓度没有明显变化，Cl^-浓度大幅度增加，而碱胁迫下有机酸浓度大幅度增加，Cl^-浓度没有明显变化.盐、碱胁迫下小冰麦茎叶中的阳离子均以Na⁺和K⁺为主，但阴离子的来源明显不同.盐胁迫下无机阴离子对负电荷的贡献起主导作用，其贡献率达61.3%～66.7%；而碱胁迫下，随胁迫强度的增大，有机酸对负离子的贡献率从38.35%上升到61.60%，逐渐成为主导成分.实验结果表明，有机酸积累是小冰麦在碱胁迫下保持体内离子平衡和pH稳定的关键生理响应.