Effects of Exogenous Glycinebetaine on Growth and Ultrastructure of Salt-Stressed Rice Seedlings (Oryza sativa L.)

Abstract

The effects of exogenously applied glycinebetaine on the salt-stress-induced inhibition of growth and ultrastructural damages in rice seedlings were investigated. Glycinebetaine was not effective in alleviating the NaCl-induced inhibition of root growth and rather enhanced the NaCl-induced inhibition. However, it was found to alleviate the inhibition of shoot growth induced by NaCl stress. Concentrations of Na were higher in salt-stressed plants than in unstressed plants. Stressed plants receiving glycinebetaine had a significantly lower Na and higher K concentrations in the shoots than the plants grown without application of glycinebetaine. Salinity induced ultrastructural damages in leaf such as swelling of thylakoids, disintegration of grana stacking and intergranal lamellae and destruction of mitochondria (deficiency of cristae, swelling and vacuolation). Such damages were largely prevented by pretreatment with glycinebetaine resulting in greening of the plants. In roots, the epidermis, cortex and root cap were more sensitive to salt stress than the meristem and stele. The most frequently observed ultrastructural alteration due to NaCl salinity was the formation of many large vacuoles in the root tip and root cap cells. The number of mitochondria was increased and they were aggregated in the cytoplasm of the root tip and root cap cells by treatment with NaCl or NaCl plus glycinebetaine. Glycinebetaine could not prevent the NaCl-induced ultrastructural damages in root cells. The effects of glycinebetaine to mitigate the ultrastructural damages in the chloroplast and mitochondria induced by NaCl might be due to the production of many vacuoles in root cells which may act to store Na and decrease its accumulation in the shoot.     

Salinity-Induced Ultrastructural Alterations in Leaf Cells of Rice (Oryza sativa L.)

Summary

High concentrations of NaCl significantly reduced the fresh and dry weights and lengths of roots and shoots. NaCl exhibited a more rapid effect in water culture than in soil culture. In both water and solid cultures, root growth was suppressed more severely than shoot growth. Electron microscopic studies revealed that NaCl caused swelling of thylakoids, accumulation of starch grains and lipid droplets, distortion of grana stacking, increase in the size and number of plastoglobuli and vesiculation of cellular membrane. Mitochondria became deficient in cristae, swelled and the matrix appeared pale in salt-treated plants as compared with control plants. Disappearance of nucleolus and nuclear chromatin and destruction of vascular tissues were occasionally observed in salt-treated plants.     

Response of Sunflowers to Quantities of Irrigation Water,Irrigation Regimes and Salinities in the Water and Soil

Summary

The production of sunflower grains for roasting was investigated in two soil types under different quantities of applied saline and non-saline irrigation water, different irrigation managements, soil salinity due to previous use of saline water or due to a raised water table. It was shown in one experiment, conducted in a loess type soil, that sunflowers extracted water at least to a soil depth of 120 cm, when the available water from the top layers was used up. The crop in this soil consumed all the available soil water from nearly the entire root zone, while in the clay soil limited water was consumed from deep layers, due to the high salinity and lack of aeration.

No decrease in yield was found in the loess soil when 75% of the full amount of water (which was 0.8 of Class A pan evaporation rate) was applied. When only 50% was applied a significant decrease in yield was obtained. In contrast, in the clay soil even 75% of the full amount of water decreased the yield remarkably. Under dry-land conditions approximately 65% of maximum yield was found in the loess soil but only 45% in the clay soil. These differences are all attributed to a shallow active root system in the clay soil. Residual soil salinity from previously use of saline water had no effect on grain production in the loess soil, while saline irrigation water applied during the irrigation season decreased production, but only when water supply was not rate limiting. The combination of saline water and residual soil salinity had a marked effect on the decrease of grain yield under limited irrigation. In both soils a reduction in the amount of water applied per single irrigation and maintaining the entire irrigation period caused a significantly smaller decrease in yield than shortening the irrigation period and applying the full demand.     

Differential Sensitivity of Rice Cultivars to Salinity and Its Relation to Ion Accumulation and Root Tip Structure

Abstract

Effects of NaCl on the growth, ion content, root cap structure and Casparian band development were examined in four rice (Oryza sativa L.) cultivars with different salt resistance (salt-sensitive indica-type IR 24 and japonica-type Nipponbare and salt-resistant indica-type Nona Bokra and Pokkali). Experiments were conducted to find the differences in salinity resistance during early seedling and developed seedling stages among the cultivars. For salinity treatment, sodium chloride (NaCl) was added to nutrient solution at concentrations of 0, 25 and 50 mM for 7 days from germination to the 7th day (early seedling stage) or from the 7th day to 14th day (developed seedling stage). Growth inhibition by salinity was more prominent in the early seedling stage than in the developed seedling stage. Based on the growth, the order of the sensitivity was IR24 > Nipponbare > Nona Bokra > Pokkali. The growth of NaCl-treated rice cultivars relative to control was significantly and negatively correlated with the Na⁺ content and Na⁺/K⁺ ratio in roots and shoots in both stages. Scanning electron microscopic observation revealed that the root cap tissues proliferated and extended to the basal part of the root tip by salinity. The length of root cap was, however, reduced by 50 mM NaCl in sensitive cultivars due to peeling off. An endodermal Casparian band was formed in the basal region of the root tip. Development of the Casparian band was more prominent in sensitive cultivars than in tolerant cultivars. Root cap proliferation might be related to NaCl resistance in rice seedlings, but the Casparian band may not function efficiently in Na⁺ exclusion. Essentially the present results suggest that exclusion of Na⁺ from roots plays a critical role in expression of Na⁺ resistance in rice seedlings and the root cap is important for Na⁺ exclusion.     

Natural silicates mixed with organic fertilizers enhance corn adaptation to salt stress and improve physical characteristics of sandy soil

Soil salinity is one of the major environmental constraints to crop productivity worldwide. Therefore, the development of cost-effective and environment-friendly techniques allowing increased crop productivity and soil fertility under saline conditions is rather urgent today. The objective of this investigation was to study the effects of mixtures containing natural silicates (analcite, bergmeal, and potassium silicate) and organic fertilizers (sapropel, peat) in corn (Zea mays L.). We specifically evaluated tolerance of corn to salinity stress and certain characteristics of saline soil (viz., redox potential, conductivity, and phytotoxicity) using a factorial pot experiment, modeling NaCl salinity levels of 0, 50, 100, 150, and 200 mM under greenhouse conditions. Growth, water balance, photosynthesis, catalase activity, and accumulation of nonenzymatic antioxidants (flavonoids and anthocyanins) were measured and evaluated. Salinity stress reduced shoot and root biomass by 8–49%, photosynthetic pigment content in leaves by 15–30%, deteriorated water balance, and activated nonspecific adaptive reactions (i.e., accumulation of enzymatic and nonenzymatic antioxidants) in the corn seedlings. All the tested silicon-containing mixtures stimulated corn seedling resistance to salt stress and reduced soil phytotoxicity. This was reflected in the stimulation of growth of the corn seedlings (accumulation of shoot biomass, and formation and growth of lateral roots). The content of photosynthetic pigments, flavonoids, anthocyanins, catalase activity increased 1.3–2 times compared with plants that received NaCl only. The difference between treatments and control was most pronounced at moderate levels of salinity (100–150 mM). The mixture containing silicon minerals and sapropel (9:1 proportion) showed the highest protective effect against salinity stress.     

Characterization of the effect of high salinity on roots of Chloris gayana Kunth: carbohydrate and lipid accumulation and growth

Branched nodal roots comprise the largest portion of the root system mass in Chloris gayana Kunth and the effects of high salinity on nodal root appearance and elongation rates were analysed in cv. Boma in greenhouse experiments. Roots from salt-treated plants (0·2 mol l^–1 NaCl) were smaller than controls, and accumulated higher concentrations of soluble sugars and reserve lipids. The number of nodal roots was reduced by the saline treatment. Leaf, tiller and nodal root appearance were delayed by salinity but the correlation among these processes was maintained, indicating that the developmental pattern was not altered by this level of salinity, only its rate. Initial nodal root growth rates varied as a function of plant size and were decreased by salinity only after 2 weeks of treatment. When shoots of non-salinized plants were cut, a drastic reduction in nodal root appearance was observed, suggesting emerging leaves were stronger sinks than roots for available reserves. However, when the shoots of salt-treated plants were cut, the already depressed root appearance rate was not further reduced. This suggests that, under salinity, alterations in root ability to metabolize reserves could have been more significant than reserve availability for controlling elongation.     

The Use of Saline Water in Agriculture in the Near East and North Africa Region: Present and Future

SUMMARY

Salinity is a major problem that negatively impacts agricultural activities in many regions in the world, and especially the Near East and North Africa region. Generally, salinity problems increase with increasing salt concentration in irrigation water. Crop growth reduction due to salinity is generally related to the osmotic potential of the root-zone soil solution. This will lead to certain phenological changes and substantial reduction in productivity. Salinity also affects the soil physical properties. Sewage treated wastewater is an alternative water source for irrigation. Using such wastewater will provide a new water resource to expand agricultural activities as well as reduce the environmental pollution. Each country in the region has a unique system of rules and regulations to protect the quality of water resources. Important aspects that should be taken into account when using wastewater for irrigation are discussed, including some information on the different irrigation systems used in the region, and the factors leading to success of using saline water for economic crop production. Information on the use of saline water or marginal saline soils for wheat production and improvement of irrigation systems, including old land irrigation systems, under Egyptian conditions is also presented. The regional experiences and the future prospects of using saline water for crop production that vary greatly among countries in the Near East and North Africa region are summarized. This article also presents information on special cultivation methods, such as protected agriculture and soilless culture that can help in alleviating the salinity effects. Finally, the article includes some examples on the inherited knowledge for saline agriculture that conveys the grower's experience in the Near East and North Africa region where several living examples for unique and sustainable cultivation system are still in operation. One of the most impressive cultivation techniques for bio-saline crop management in Egypt is the Edkawy production system.     

Vertical Distribution of Sodium in Roots of Rice Plants Exposed to Salinity as Analyzed by Cryo Time-of-Flight Secondary Ion Mass Spectrometry

Abstract

Distribution of Na⁺ along the root axis under salinity stress was analyzed in two rice (Oryza sativa L.) cultivars with different salt resistance (salt-sensitive IR 24 and salt-resistant Pokkali). Rice plants were grown hydroponically and NaCl was applied with nutrient solution at concentrations of 0, 25 and 50 mM for 7 d after germination. The distribution of Na⁺ in roots under salinity was analyzed by the cryo time-of-flight secondary ion mass spectrometry (cryo TOF-SIMS). The Na⁺ content in the root was higher in salt-sensitive IR 24 than in salt-resistant Pokkali under NaCl stress. The content was highest at the root tip and was decreased basipetally along the root axis. The difference in Na⁺ content between the cultivars was apparent in all regions from the root tip.     

Effect of saline soil on grain quality of rices differing in salinity tolerance

Four varieties of rice, differing in salinity tolerance and grown in saline soil (electrical conductivity 5–6 dS/m) at Sadhoke, Punjab, Pakistan, had lighter grain and higher Na content than control samples. Grains of three out of the four rices grown on saline soils had higher brown rice protein (higher nutritional value), less translucent grain, lower starch and amylose content, and lower K than their control samples, but these differences were not related to salinity tolerance. Alkali spreading value and gel consistency were not affected by culture in saline soil. Cooked rice Instron hardness increased in saline culture in two higher-protein samples of the four rices. Amylograph peak viscosity was suppressed by saline culture.     

In vitro bioassays for salinity tolerance screening of potato

Summary Three in vitro bioassays (single-node cutting, root tip segment and microtuberization) gave similar results in ranking the salinity (NaCl) tolerance of several potato genotypes (Solanum spp.) and were verified by tuber yield criteria in a field lysimeter trial with salinized irrigation water. Salinity stress consistently depressed growth in the single-node cutting and root tip segment bioassays, reduced microtuber yield in the microtuberization bioassay and decreased tuber yield in field lysimeters. The single-node cutting bioassay was simpler to perform than the root tip segment and microtuberization bioassays and did not exclude certain genotypes as did the microtuberization bioassay. The single-node cutting bioassay can be recommended as a substitute for more labour-intensive and costly field assessments of salinity effects of yield.     

盐胁迫对垂盆草生长和生理特性的影响

以垂盆草为材料,研究不同浓度NaCl[0(对照)、25、50、100、150、200 mmol/L]对垂盆草生长和生理特性的影响.结果表明:与对照相比,随着NaC1浓度的增加,垂盆草鲜重、干重、含水量、K+含量、根Na+/叶Na+、叶的K+/Na+和SOD活性均呈下降趋势;而Na+、游离氨基酸、可溶性糖、有机酸、脯氨酸、丙二醛含量和质膜透性均呈上升趋势;叶绿素含量、硝酸还原酶活性呈先上升后下降趋势,而NO3-含量则呈相反的变化趋势.综合分析显示,盐胁迫对垂盆草生长有抑制作用,其耐盐阈值为100 mmol/L NaCl;盐胁迫下垂盆草通过积累Na+、有机酸、游离氨基酸、可溶性糖、脯氨酸进行渗透调节.     

Effect of Azotobacter vinelandii and compatible solutes on germination wheat seeds and root concentrations of sodium and potassium under salt stress

The effect of plant growth-promoting Rhizobacteria (PGPR) and exogenous application of compatible solutes on seed germination and root concentrations of sodium and potassium of two wheat varieties (Triticum durum L.) were evaluated under saline stress. In this experiment, Azotobacter vinelandii strain DSM85, glycine betaine and proline were used. Inoculated seeds for each variety were placed on Whatman paper in 9 cm Petri dishes containing 15 mL of distilled water or NaCl solutions at various concentrations (control, 100, 200, 300 mM) supplemented with or without glycine betaine (GB) or proline at 5 mM. The results indicated that addition of proline (5 mM) stimulated the production of indol acetic acid and the growth of A. vinelandii at 200 and 300 mM NaCl, respectively. The germination rate index and the germination final percentage decreased significantly (p < 0.05) with increasing salinity level. The germination was significantly diminished at 300 mM with significant variation among varieties and Waha variety had higher germination percentage than Bousselam variety. Inoculation of seeds by A. vinelandii and exogenous application of proline had significantly positive effect on the germination at this concentration of NaCl. The rate of accumulation of Na+ in roots was important at 100 mM and increased at 200 mM. The concentration of K+ decreased when salinity increased. The effect of inoculation or inoculation with proline decreased the accumulation of Na' and reduced the loss of K+ under salt stress. From the present study we can conclude that the use of A. vinelandii strain DSM85 and external application of low concentrations of proline on seeds might be considered as a strategy for the protection of plants under saline stress.     

Long-Term Reuse of Drainage Waters of Varying Salinities for Crop Irrigation in a Cotton-Safflower Rotation System in the San Joaquin Valley of California—A Nine Year Study: I. Cotton (Gossypium hirsutum L.)

Summary

Use of saline drainage water for crop irrigation was evaluated as a means of decreasing its volume. Results of a nine-year crop rotation (cotton-cotton-safflower, × 3) in which only the cotton was irrigated with drainage water of 400, 1,500, 3,000, 4,500, 6,000, and 9,000 ppm total dissolved salts are presented. The different salinity levels of irrigation waters were achieved by mixing nonsaline canal water (400 ppm) and saline drainage water. Cotton lint yields were not affected by increased salinity level of the irrigation water for the first two years. Detrimental effects became evident in the third cotton crop with increasing severity in later years. In the fifth year of cotton (seventh year of the study), lint yields were adversely affected by waters of salinity greater than 3,000 ppm. However, fiber quality remained unaffected at all levels of irrigation water salinity. The reductions in lint yield appeared to be a function of time and the salinity level of applied water. Shoot height and biomass were reduced by the irrigation water salinity before lint yields. Stand establishment appeared to be the most sensitive to salinity and was perhaps the main reason for yield reduction. Increase in irrigation water salinity increased Na⁺ content of leaf blades and petioles and decreased K⁺/Na⁺ ratio of leaf blades and petioles. The study showed that irrigation waters of up to 3,000 ppm salinity may be used for four years without any yield reductions, as long as some leaching occurs through preplant irrigations with low salinity water. Data on crop growth and development and ionic content collected over the nine year period are presented.     

Differential Sensitivity of Chloroplasts in Mesophyll and Bundle Sheath Cells in Maize,an NADP-Malic Enzyme-Type C4 Plant,to Salinity Stress

Abstract

The changes in chloroplast ultrastructure and the contents of chlorophyll, Na and K in response to salinity stress were investigated in leaves of maize, an NADP-malic enzyme-type C₄ plant species possessing dimorphic chloroplasts. The seedlings were treated with 0, 1, 2 or 3% NaCl for three or five days under a light or dark condition. In both light and dark conditions, the dry weight of salt-treated plants decreased as NaCl concentration increased. Chlorophyll and K contents of the second leaf blade decreased as NaCl concentration increased under the light condition but not under the dark condition. Na content of the second leaf blade was significantly higher at high NaCl concentrations under both light and dark conditions. However, Na content was much lower under the dark condition than light condition. Higher concentrations (2 and 3%) of NaCl significantly increased the size of plastoglobules, decreased the number and size of starch granules and altered the chloroplast ultrastructure. Under the light condition, mesophyll cell (MC) chloroplasts appeared more sensitive to the damaging effect of salinity than the bundle sheath cell (BSC) chloroplasts. MC chloroplasts became more globular in shape and showed swollen and disorganized thylakoids and reduced thickness of grana by salinity. BSC chloroplasts were less affected by salinity than MC chloroplasts. Although chloroplast size and number and size of starch granules were reduced, there was no structural distortion in the thylakoids of BSC chloroplasts. However, the thickness of grana was increased by salinity. Under the dark condition, the chloroplast structure was less affected by salinity. Though the envelope of BSC chloroplasts was occasionally damaged, the thylakoids in both MC and BSC chloroplasts were preserved under salinity stress. The present study suggests that the chloroplast damage caused by salinity is light-dependent and MC chloroplasts are more sensitive to salinity than BSC chloroplasts.     

混合盐碱胁迫对高粱幼苗的影响

用50，100，200，300mmol/L含有不同比例NaCl,Na2SO4,NaHCO3和Na2CO3的混合盐溶液对高粱（Sorghum bicolor L.）幼苗进行混合盐碱胁迫处理。测定高粱幼苗存活率、相对生长率、相对含水量、相对根系活力以及电解质外渗率等胁变指标。结果表明：上述各项胁变反应均随盐浓度和碱性盐比例增加而加剧，由碱性盐造成的高pH所致的胁变效应与盐浓度之间相关：即低盐度时pH作用较小，随盐度增大其作用加剧。在碱胁迫较弱时胁变主要受盐度影响，随碱胁迫增大pH成为影响胁变的主要因素。在高盐高碱条件下，盐胁迫与碱胁迫间具有协同作用。     

Regulation of root-to-leaf Na and Cl transport and its association with photosynthetic activity in salt-tolerant soybean genotypes

ABSTRACT

Soil salinity is a major constraint to sustainable crop production. Genetic improvements are needed for growing soybean in salinity-prone environments. Salt-tolerant soybean genotypes alleviate a reduction in photosynthesis and growth under saline conditions; however, the detailed mechanisms involved remain unclear. Here, we aimed to clarify how Na and Cl root-to-leaf transport is quantitatively regulated, and to identify whether photosynthetic tolerance depends on traits associated with either stomata or with mesophyll tissues. Two pairs of pot-grown soybean near-isogenic lines (NILs) consisting of tolerant and susceptible counterparts, derived from a cross between salt-tolerant FT-Abyara and salt-sensitive C01, were subjected to salinity treatment in a rainout greenhouse. Comparison of photosynthetic responses between genotypes indicated that genotypic differences in salinity tolerance depended on the ability for sustained CO₂ assimilation in mesophyll tissues, rather than stomatal conductance. The ratio of photosynthetic rate to intercellular CO₂ concentration (A/Ci) declined exponentially with increasing Na and Cl concentration regardless of genotype, but tolerant genotypes effectively kept both elements at significantly low levels. Under saline conditions, tolerant genotypes reduced Na and Cl content at the two transport pathways: from root to stem, and from stem to leaf, but the reduction of Cl at each pathway was only minor. These results suggest that integrating genetic capacity for Cl transport regulation and osmotic adjustment should be an important target in salinity-tolerance soybean breeding.     

Molecular Approaches to Improve Salt Resistance in Crops: Facts and Perspectives

SUMMARY

Because of the expansion of agriculture into marginal environments, enhancement of crop resistance to soil salinity is becoming a frequent objective for breeders. The tools offered by molecular biology to transfer a single or a few genes provide a major hope to reduce the negative impact of broad gene transfer that takes place in wide-cross hybridizations. Due to the presence of osmotic and toxic components in the growth response of plants to salt stress, any attempt to improve plant performance in saline environments should ensure the maintenance of an adequate flux of water into plant tissues, and also avoid the build up of ions into the cell compartments where they can exert toxic effects. Besides, reduction of injury effects due to salinity on plant tissues is a highly desirable objective. Transgenic plants overexpressing ion transporters able to exclude Na⁺ into vacuoles, the enzymes required for the biosynthesis of several osmocompatible, organic solutes, or the enzymes participating in detoxification pathways, have been obtained. Some of these transgenic plants display an enhanced growth relative to their wild type parents in saline environments, although the way in which this resistance is achieved remains essentially unknown. A fourth and promising way to engineer salt resistance in plants is the attempt to manipulate gene regulatory pathways. The extent to which these experiences, mainly with model plants, could be extrapolated to crop plants growing in the field is discussed. It is proposed that a combination of different molecular approaches could be helpful to achieve enhanced salt resistance in crop plants.     

Long-Term Reuse of Drainage Waters of Varying Salinities for Crop Irrigation in a Cotton-Safflower Rotation System in the San Joaquin Valley of California—A Nine Year Study: II. Safflower (Carthamus tinctorius L.)

Summary

This paper reports the results on safflower crops grown in a nine-year study, conducted on a 8 ha site, to determine the feasibility of using drainage water for irrigation in a 2-year cotton/1-year safflow-er rotation system. The cotton crops were irrigated with waters of 400, 1,500, 3,000, 4,500, 6,000, and 9,000 ppm total dissolved salts, and safflower was grown only with a preplant irrigation with nonsaline water. The use of drainage water for crop irrigation may be a means of decreasing its volume. Even though safflower was never irrigated with saline drainage water directly, the residual effect of using saline water for cotton irrigation adversely impacted safflower growth and development. Safflower seed yields were reduced in plots previously irrigated with waters of 4,500 ppm or higher salinity and even more severe effects on crop growth were seen as the numbers of years of irrigation with the saline water increased. After irrigating six cotton crops, the safflower seed yield in plots irrigated with 9,000 ppm waters was reduced to only 14% of the control. The safflower oil content and quality were not affected. Impacts on plant density, shoot height, shoot biomass, and leaf ionic content also are discussed.     

Field Crop Production in Areas with Saline Soils and Shallow Saline Groundwater in the San Joaquin Valley of California

SUMMARY

Salinity in soil and water is irrevocably associated with irrigated agriculture throughout the world and as a result requires that salt management becomes an integral part of the production system. With careful water management, it is possible to sustain irrigated agriculture in areas with saline soil and saline groundwater with and without subsurface drainage. The results from two field projects conducted in an area with saline soils and saline groundwater demonstrated the type of irrigation systems and management needed to sustain production of moderately salt tolerant and tolerant crops. During the first study at Murrieta farms, yields of cotton and sugar beet were maintained using both saline and non-saline water for irrigation when pre-plant irrigation and rainfall were adequate to maintain soil salinity at a tolerable level. Wheat production was reduced in areas that used saline water for irrigation. Use of saline water containing toxic elements such as boron for irrigation poses a threat to the sustainability of the system. The second study evaluated the management of furrow and subsurface drip irrigation in the presence of shallow saline groundwater. Careful management of the furrow system during pre-plant irrigation and the first irrigation of the growing season was required to prevent waterlogging. It was possible to manage the subsurface drip system to induce significant crop water use from shallow groundwater. Rainfall and pre-plant irrigation were adequate at this site to manage soil salinity.     

Genetic Opportunities to Improve Cereal Root Systems for Dryland Agriculture

Abstract

Understanding the major limitations to root growth is very important if we are to maximize water and nutrient use and increase yields. Limitations may be insufficient rooting depth, root diseases, nutrient deficiencies, toxicities and soil hardness. An understanding of these limitations will lead to more precisely identifying traits for which to select and breed. Examples of successfully overcoming limiting factors to improve crop performance by breeding and selection are given for cereal cyst nematodes in wheat, soil acidity and salinity. The importance of altered crop management practices to reduce limitations is also stressed. These have resulted in a more effective and healthier root system, which results in more water use and greater yields. Opportunities to genetically increase the size of the root system in dryland systems where water and nutrients are not all used by the crop are given.