Anatomical adaptations to salinity in cogon grass [Imperata cylindrica (L.) Raeuschel] from the Salt Range, Pakistan

To examine anatomical adaptations in a potential forage grass, Imperata cylindrica (L.) Raeuschel, a population was collected from the natural salt-affected soils of the Salt Range, Pakistan. Using a hydroponic system, the degree of salt tolerance in terms of structural modifications in the Salt Range ecotype was compared with that in an ecotype collected from a normal non-saline habitat of the Faisalabad region. The Salt Range ecotype was superior to the Faisalabad ecotype in biomass production under saline conditions. High salt tolerance of the Salt Range ecotype was associated with increased succulence in root and leaf (mainly midrib), formation of aerenchyma in leaf sheath, increased vascular bundle area, metaxylem area and phloem area, highly developed bulliform cells on leaves and increased sclerification in root and leaf. Furthermore, both stomatal density and stomatal area were considerably reduced under high salinities in the Salt Range ecotype.     

Physiological and biochemical adaptations of Cynodon dactylon (L.) Pers. from the Salt Range (Pakistan) to salinity stress

Naturally adapted salt tolerant populations provide a valuable material for exploring the adaptive components of salt tolerance. Under this aspect, two populations of Cynodon dactylon (L.) Pers. were subjected to salt stress in hydroponics. One was collected from a heavily salt-affected soil in the vicinity of a natural salt lake, Uchhali Lake, in the Salt Range of the Punjab province of Pakistan, and the other from a normal non-saline habitat from the Faisalabad region. The NaCl treatments in Hoagland's nutrient solution were: Control (no salt), 50, 100, 150 and 200 mM of NaCl. After 8 weeks of growth in hydroponics produced biomass, ion relations, and photosynthetic capacity were measured in the differently adapted ecotypes. In the ecotype of C. dactylon from the Salt Range, shoot dry weight was only slightly affected by varying levels of salt. However, in contrast, its root weight was markedly increased. On the other hand, the ecotype from Faisalabad (non-saline habitat) showed a marked decrease in shoot and root dry weights under saline regimes. The ecotype from the Salt Range accumulated relatively less amount of Na⁺ in the shoot than did that from Faisalabad, particularly at higher salt levels. Shoot or root K⁺ and Ca²⁺ contents varied inconsistently in both ecotypes under salt stress. All the photosynthetic parameters, leaf water potential and osmotic potential, and chlorophyll content in both ecotypes were adversely affected by salt stress, but all these physiological attributes except turgor potential and soluble sugars were less affected at high salinities in the salt tolerant ecotype from Salt Range. This ecotype accumulated significantly higher organic osmotica (total free amino acids, proline, total soluble proteins, and total soluble sugars) under saline conditions than its intolerant counterpart. Overall, the salt tolerant ecotype of C. dactylon from the Salt Range showed high salt tolerance due to its restricted uptake of Na⁺ accompanied by an increased uptake of K⁺ and Ca²⁺ in the roots as well as shoot due to its higher photosynthetic capacity and accumulation of organic osmotica such as free amino acids and proline under saline conditions.     

Physiological adaptative characteristics of Imperata cylindrica for salinity tolerance

Two populations of cogongrass [Imperata cylindrica (L.) Raeuschel], one from the saline regions of the Salt Range and the other from the non-saline regions of Faisalabad were assessed for salinity tolerance on the basis of some key morphological and physiological attributes. It was hypothesized that the tolerant population from the Salt Range must have developed some specific structural modifications, which are responsible for its better survival under high salinities. These adaptive components can be effectively used in modern technologies for improving degree of tolerance of other sensitive crops. The population from the Salt Range markedly excelled the Faisalabad population in terms of growth and physiological attributes measured in this study. The Faisalabad population of I. cylindrica was unable to survive at the highest salt level (200 mM NaCl). The tolerance of the Salt Range population to salt stress was found to be related to high accumulation of organic osmotica, particularly total free amino acids and proline as well as Ca²⁺ in the shoot. The distinctive structural modifications in the Salt Range population were found to be enhanced succulence, well-developed bulliform cells in leaves and smaller stomatal area.     

Control of xylem Na+ loading and transport to the shoot in rice and barley as a determinant of differential salinity stress tolerance

Control of xylem Na⁺ loading has often been named as the essential component of salinity tolerance mechanism. However, it is less clear to what extent the difference in this trait may determine differential salinity tolerance between species. In this study, barley (Hordeum vulgare L. cv. CM72) and rice (Oryza sativa L. cv. Dongjin) plants were grown under two levels of salinity. Na⁺ and K⁺ concentrations in the xylem sap, and shoot and root tissues were measured at different time points after stress onset. Salt‐exposed rice plants prevented xylem Na⁺ loading for several days, but failed to control this process in the longer term, ultimately resulting in a massive Na⁺ shoot loading. Barley plants quickly increased xylem Na⁺ concentration and its delivery to the shoot (most likely for the purpose of osmotic adjustment) but were able to reduce this process later on, keeping most of accumulated Na⁺ in the root, thus maintaining non‐toxic shoot Na⁺ level. Rice plants increased shoot K⁺ concentration, while barley plants maintained higher root K⁺ concentration. Control of xylem Na⁺ loading is remarkably different between rice and barley; this difference may differentiate the extent of the salinity tolerance between species. This trait should be investigated in more detail to be used in the breeding programs aimed to improve salinity tolerance in crops.     

Salt Tolerance ofEchinochloa crusgalli

Salt tolerance ofEchinochloa crusgalli was studied using gravel culture with root medium electrical conductivity between 3 to 25 dS m^-1. Salinity depressed germination and shoot yield. A 50 % reduction in shoot yield occurred at 15.9 dS m^-1. The plant was able to maintain its tissue water content and K concentration in the tissue water while Na, Ca and Cl increased and Mg decreased with increasing root zone salinity.     

Exogenously applied paclobutrazol modulates growth in salt-stressed wheat plants

Effect of paclobutrazol (PBZ) treatment on salinity tolerance of wheat (Triticum aestivum) was investigated on a salt-tolerant (Karchia-65) and salt-sensitive (Ghods) cultivars. Salinity significantly reduced the investigated growth parameters such as plant height, length and area of sixth leaf, root length, fresh and dry weight of shoot, roots and sixth leaf, water content (WC) of plant and seeds weight in the both cultivars. The negative effect of salinity in Ghods cultivar was more than Karchia cultivar. However, PBZ treatment reduced the growth in both cultivars, the differences in plant growth among various levels of NaCl decreased in PBZ-treated plants. Salt stress resulted in high accumulation of Na⁺ in the sixth leaf and roots in both cultivars, particularly in Ghods cultivar. Against Karchia cultivar, salt stress decreased the storage of K⁺, P and N in sixth leaf and roots in Ghods cultivar. In the both cultivars, PBZ treatment enhanced the K⁺, P and N contents in sixth leaf and roots by increasing salinity. Although PBZ treatment decreased the growth of plants, it improved the weight of seeds against stress damage. PBZ treatment reduced the accumulation of harmful Na⁺ ion in plant tissues while increased the K⁺, P and N contents. These observations suggest that PBZ treatment may increase tolerance by diminishing ionic imbalance caused by salt stress.     

Identification of quantitative trait loci for ion homeostasis and salt tolerance in barley (Hordeum vulgare L.)

Ion homeostasis is considered to be one of the most important mechanisms underlying salt stress tolerance. We used the Steptoe × Morex barley doubled haploid population to screen for genetic variation in response to salinity stress at an early development stage in a hydroponics system, focusing on ion homeostasis. Salinity induced a strong adverse effect on growth of the parents and their derived population, with Steptoe as the more tolerant parent. Steptoe maintained higher concentrations of K⁺, Na⁺ and Cl^? in the roots and a similar shoot/root ion ratio (<1) under stress conditions compared to control conditions. In contrast, Morex had higher concentrations of these ions in the shoots under stress and a doubled shoot/root ion ratio relative to control conditions, indicating that salt exclusion might contribute to the higher tolerance of Steptoe. Correlation and path analysis demonstrated that shoot Cl^? contents most strongly affected salt tolerance and suggest that both Na⁺ and Cl^? contents are important for salinity stress tolerance in barley. We identified 11 chromosomal regions involved in the control of the variation observed for salt tolerance and various salt stress response traits, including Na⁺, Cl^? and K⁺ contents in shoots. Two specific regions on chromosomes 2H and 3H were found controlling ion contents and salt tolerance, pointing to genes involved in ion homeostasis that contribute to salt tolerance.     

A single nucleotide substitution in TaHKT1;5-D controls shoot Na+ accumulation in bread wheat

Improving salinity tolerance in the most widely cultivated cereal, bread wheat (Triticum aestivum L.), is essential to increase grain yields on saline agricultural lands. A Portuguese landrace, Mocho de Espiga Branca accumulates up to sixfold greater leaf and sheath sodium (Na⁺) than two Australian cultivars, Gladius and Scout, under salt stress in hydroponics. Despite high leaf and sheath Na⁺ concentrations, Mocho de Espiga Branca maintained similar salinity tolerance compared to Gladius and Scout. A naturally occurring single nucleotide substitution was identified in the gene encoding a major Na⁺ transporter TaHKT1;5-D in Mocho de Espiga Branca, which resulted in a L190P amino acid residue variation. This variant prevents Mocho de Espiga Branca from retrieving Na⁺ from the root xylem leading to a high shoot Na⁺ concentration. The identification of the tissue-tolerant Mocho de Espiga Branca will accelerate the development of more elite salt-tolerant bread wheat cultivars.     

Morphological responses of two plant species from different elevations in the Dongting Lake wetlands,China, to variation in water levels

The vegetation of wetlands show strong zonation patterns, but the mechanisms determining these patterns are not fully understood. In the present study, growth and morphological responses to a water level gradient (–20 cm (i.e. water level 20 cm below soil surface), –10 cm, 0 cm, 10 cm, 20 cm) were compared between a higher elevation plant (Imperata cylindrica) and a lower elevation plant (Carex brevicuspis) in the Dongting Lake wetlands of China. For both species, the aboveground, belowground, and total biomass were greater at –10 cm than at any other water level.. However, when the water level increased from –10 cm to 0 cm, there was a greater decrease in the biomass of I. cylindrica than in that of C. brevicuspis. Plant height, tiller number, leaf length, leaf width and leaf area showed greater variation along the water level gradient in I. cylindrica than in C. brevicuspis. Generally, with increasing water level, root length, rhizome number, and adventitious root biomass and number all decreased in I. cylindrica. However, in C. brevicuspis, neither the rhizome number nor the primary adventitious root biomass differed significantly among the five water levels. These results indicate that I. cylindrica have a lower tolerance for flooding and higher water sensitivity than C. brevicuspis and these differences may explain why I. cylindrica is found at relatively higher elevations that are not prone to flooding, while C. brevicuspis is found at comparatively lower elevations in the Dongting Lake wetlands.     

Triacontanol modulates photosynthesis and osmoprotectants in canola (Brassica napus L.) under saline stress

Abstract

An experiment was conducted to assess the effect of pre-sowing seed treatment with triacontanol (TRIA) in canola (Brassica napus L.) cultivar (RBN-3060) under saline stress. Canola seeds were soaked in three levels of TRIA (0, 0.5, and 1 mg L^?1) for 12 hours. Three levels of salt stress (0, 100, and 150 mM NaCl) in full strength Hoagland's nutrient solution were applied to 56-days-old plants. Salt stress caused a significant reduction in growth, gas exchange, photochemical quenching (qP), and shoot and root K⁺ contents, while increased leaf glycine betaine, free proline, and shoot Na⁺ contents. Pre-sowing seed treatment with TRIA increased shoot fresh weight, number of seeds per plant, photosynthetic rate, transpiration rate, ratio of chlorophyll a/b, qP, electron transport rate, shoot and root K⁺ contents, and free proline and glycine betaine contents of canola plants at various TRIA levels under nonsaline or saline conditions.     

Association mapping of salt tolerance in barley (Hordeum vulgare L.)

A spring barley collection of 192 genotypes from a wide geographical range was used to identify quantitative trait loci (QTLs) for salt tolerance traits by means of an association mapping approach using a thousand SNP marker set. Linkage disequilibrium (LD) decay was found with marker distances spanning 2–8 cM depending on the methods used to account for population structure and genetic relatedness between genotypes. The association panel showed large variation for traits that were highly heritable under salt stress, including biomass production, chlorophyll content, plant height, tiller number, leaf senescence and shoot Na⁺, shoot Cl^? and shoot, root Na⁺/K⁺ contents. The significant correlations between these traits and salt tolerance (defined as the biomass produced under salt stress relative to the biomass produced under control conditions) indicate that these traits contribute to (components of) salt tolerance. Association mapping was performed using several methods to account for population structure and minimize false-positive associations. This resulted in the identification of a number of genomic regions that strongly influenced salt tolerance and ion homeostasis, with a major QTL controlling salt tolerance on chromosome 6H, and a strong QTL for ion contents on chromosome 4H.     

Salicylic Acid Confers Salt Tolerance in Giant Juncao Through Modulation of Redox Homeostasis,Ionic Flux,and Bioactive Compounds: An Ionomics and Metabolomic Perspective of Induced Tolerance Responses

We investigated the stimulatory and/or inhibitory role of exogenous SA in alleviating the salt stress (250, 500 mM NaCl) in Pennisetum giganteum (Giant Juncao) through coordinated induction of redox homeostasis, ionic flux, and bioactive compounds. Salt stress radically impaired root and shoot (growth, fresh, and dry biomass as well as tolerance indices), leaf relative water content, and leaf chlorophyll a/b ratio of Juncao due to higher Na⁺ and Cl^? accumulation followed by H₂O₂ generation, lipid peroxidation (MDA contents), and electrolyte leakage. However, the innate defense response of Juncao counteracted salt-induced damages by osmolytes accumulation combined with orchestrating antioxidants and ionic homeostasis mechanisms. Furthermore, the application of SA had an incremental impact on the development and productivity of high-salinity-exposed Juncao plants by increasing root length, plant biomass, tolerance indices, chlorophyll a/b ratio, and protein contents. Furthermore, SA treatment considerably decreased Na⁺ and Cl^? toxicity by orchestrating antioxidant enzymes, ion transport, and secondary metabolism. Notably, the application of SA substantially mitigated the adverse effects of high salinity concentration (500 mM NaCl), owing to the simultaneous upregulation in enzymatic and non-enzymatic antioxidants, nutrient ion flux, alongside chlorogenic acid production. Thus, we concluded that SA enhanced the tolerance capability of Juncao plants in a NaCl concentration-dependent manner. The findings of this study will enable environmentalists and pharmacologists to gain dual farm-level benefits, including animal therapeutics and restoration of salinized soils for arable purposes.

     

Salt stress response in tomato beyond the salinity tolerance threshold

Crop salt tolerance is generally assessed as the relative yield response to increasing root zone salinity, expressed as soil (EC_e) or irrigation water (EC_w) electrical conductivity. Alternatively, the dynamic process of salt accumulation into the shoot relative to the shoot biomass has also been considered as a tolerance index. These relationships are graphically represented by two intersecting linear regions, which identify (1) a specific threshold tolerance, at which yield begins to decrease, and (2) a declining region, which defines the yield reduction rate. Although the salinity threshold is intuitively a critical parameter for establishing plant salt tolerance, we focused our interest on physiological modifications that may occur in the plant at salinity higher than the so-called tolerance threshold. For this purpose, we exposed hydroponically grown tomato plants to eight different salinity levels (EC = 2.5 (non-salinized control); 4.2; 6.0; 7.8; 9.6; 11.4; 13.2; 15.0 dS m⁻¹). Based on biomass production, water relations, leaf ions accumulation, leaf and root abscisic acid and stomatal conductance measurements, we were able to identify a specific EC value (approximately 9.6 dS m⁻¹) at which a sharp increase of the shoot and root ABA levels coincided with (1) a decreased sensitivity of stomatal response to ABA; (2) a different partitioning of Na⁺ ions between young and mature leaves; (3) a remarkable increase of the root-to-shoot ratio. The specificity and functional significance of this response in salt stress adaptation is discussed.     

Enhanced formation of aerenchyma and induction of a barrier to radial oxygen loss in adventitious roots of Zea nicaraguensis contribute to its waterlogging tolerance as compared with maize (Zea mays ssp. mays)

Enhancement of oxygen transport from shoot to root tip by the formation of aerenchyma and also a barrier to radial oxygen loss (ROL) in roots is common in waterlogging‐tolerant plants. Zea nicaraguensis (teosinte), a wild relative of maize (Zea mays ssp. mays), grows in waterlogged soils. We investigated the formation of aerenchyma and ROL barrier induction in roots of Z. nicaraguensis, in comparison with roots of maize (inbred line Mi29), in a pot soil system and in hydroponics. Furthermore, depositions of suberin in the exodermis/hypodermis and lignin in the epidermis of adventitious roots of Z. nicaraguensis and maize grown in aerated or stagnant deoxygenated nutrient solution were studied. Growth of maize was more adversely affected by low oxygen in the root zone (waterlogged soil or stagnant deoxygenated nutrient solution) compared with Z. nicaraguensis. In stagnant deoxygenated solution, Z. nicaraguensis was superior to maize in transporting oxygen from shoot base to root tip due to formation of larger aerenchyma and a stronger barrier to ROL in adventitious roots. The relationships between the ROL barrier formation and suberin and lignin depositions in roots are discussed. The ROL barrier, in addition to aerenchyma, would contribute to the waterlogging tolerance of Z. nicaraguensis.     

Growth promotion effect of Fusarium spp. PPF1 from bermudagrass (Cynodon dactylon) rhizosphere on Indian spinach (Basella alba) seedlings are linked to root colonisation

A rhizosphere fungus was isolated from roots of bermudagrass (Cynodon dactylon) and identified as Fusarium spp. PPF1. A pot experiment was conducted to test its ability to promote the vegetative growth of Indian spinach seedlings (Basella alba). Indian spinach seeds were sown in sterilised field soil amended with wheat grain inoculum of PPF1 at the rate of 0.5 and 1.0% w/w, and plants were grown for 21?days in a net house. Significantly, higher germination percentage and vigour index were observed due to application of PPF1 in the potting soil. Treatment with PPF1 also significantly increased shoot length, shoot fresh weight, shoot dry weight, root length, root fresh weight, root dry weight, leaf area and leaf chlorophyll content of cucumber plants compared to non-treated control. The growth promotion rate increased with the increasing concentration of inoculum of PPF1 applied to the soil. The fungus was re-isolated from the roots of cucumber plants at higher frequencies, while a positive co-relation was found between the root colonisation ability and the plant growth enhancement by the isolate. These results suggest that growth promotion effect of Fusarium spp. PPF1 on Indian spinach (B. alba) are linked to root colonisation ability of the fungus.     

Morpho-anatomical and physiological attributes for salt tolerance in sewan grass (Lasiurus scindicus Henr.) from Cholistan Desert,Pakistan

Three differently adapted populations of sewan grass (Lasiurus scindicus Henr.) were evaluated for structural and functional adaptations to high salinity. The habitats were Derawar Fort (DF, least saline, ECe 15.21), Bailahwala Dahar (BD, moderately saline, ECe 27.56 dS m^?1) and Ladam Sir (LS, highly saline, ECe 39.18 dS m^?1) from within the Cholistan Desert. The adaptive components of salt tolerance in sewan grass were assessed by determining various morpho–anatomical and physiological attributes. The degree of salt tolerance of all three ecotypes of L. scindicus from the saline habitats was compared in a controlled hydroponic system to evaluate the adaptive components that are expected to be genetically fixed during a long evolutionary process. Salinity tolerance in the most tolerant LS population relied on increased root length and total leaf area, restricted uptake of toxic Cl^?, increased uptake of Ca²⁺, high excretion of Na⁺, accumulation of organic osmolytes, high water use efficiency, increased root, thicker leaf and cortical region, intensive sclerification, large metaxylem vessels, and dense pubescence on abaxial leaf surface. The BD population (from moderately saline soil) relied on high Ca²⁺ uptake, Na⁺ excretion, epidermal thickness, large cortical cells, thick endodermis and large vascular tissue. The DF population (from less saline soil) showed a significant decrease in all morphological characteristics; however, it accumulated organic osmolytes for its survival under high salinities. Structural modifications in all three populations were crucial for checking undue water loss under physiological stress that is caused by high amounts of soluble salts in the soil.     

Salicylic Acid Induced Salinity Tolerance Through Manipulation of Ion Distribution Rather than Ion Accumulation

In this research, the effect of different SA concentrations (0, 0.5, 1.0, 1.5, and 2.0 mM) on biological and grain yield as well as Na⁺, K⁺, Cl^?, Ca²⁺, and Mg²⁺ distribution and accumulation in barley plants was examined under nonsaline (2 dS m^?1) and saline (12 dS m^?1) conditions in a three-year field study (2012–2015 growing seasons). Storage factor (SF) was defined as the concentration of an ion in the root, as a proportion of total uptake of that ion, to quantify ion partitioning between root and shoot. Salt stress decreased SF for K⁺, Ca²⁺, and Mg²⁺ and enhanced it for Na⁺ and Cl^?, which led to reduce grain and biological yield. Nonetheless, foliar-applied SA in varying concentrations could lower some of these adverse effects on ion transport and accumulation. At the 2nd and 3rd years, unfavorable climatic conditions such as less precipitation and higher temperature intensified salt stress and decreased the alleviating impact of SA. Foliar application of SA at higher levels increased SF for Na⁺ and Cl^? ions and decreased that for K⁺ indicating that SA helped barley plants keep more Na⁺ and Cl^? and less K⁺ ions in the root system, which suggested the probable role of SA in altering ion transport within the plant in favor of salt stress tolerance. SF was found to be more correlated with grain yield under both nonsaline and saline conditions. Overall, SF might be considered as a potential criterion for salt tolerance in barley plants.     

三种水库消落带草本植物对完全水淹的适应机制研究

为了解虉草(Phalaris arundinacea)、牛鞭草(Hemarthria altissima)和狗牙根(Cynodon dactylon)3种水库消落带草本植物在完全水淹条件下的生理生态适应机制,对这3种植物的生态指标(枝条、叶和生物量)和生理指标(可溶性糖、淀粉和根系活力)的动态变化特征进行了研究。结果表明：在完全水淹条件下,虉草、牛鞭草和狗牙根都不产生新的枝条,它们通过减少总叶数、总叶长和生物量的方式来适应水淹环境。虉草和狗牙根通过减缓枝条生长速率来适应水淹环境,而牛鞭草则是通过先加快生长后抑制生长来保存活力。虉草、牛鞭草和狗牙根均以少量的碳水化合物(可溶性糖和淀粉)消耗,降低根系活力等方式来适应长期的水淹环境。3种草本植物的水淹耐受能力由大到小依次为狗牙根、虉草、牛鞭草。     

A teosinte-derived allele of an HKT1 family sodium transporter improves salt tolerance in maize

The sodium cation (Na⁺) is the predominant cation with deleterious effects on crops in salt-affected agricultural areas. Salt tolerance of crop can be improved by increasing shoot Na⁺ exclusion. Therefore, it is crucial to identify and use genetic variants of various crops that promote shoot Na⁺ exclusion. Here, we show that a HKT1 family gene ZmNC3 (Zea mays L. Na⁺ Content 3; designated ZmHKT1;2) confers natural variability in shoot-Na⁺ accumulation and salt tolerance in maize. ZmHKT1;2 encodes a Na⁺-preferential transporter localized in the plasma membrane, which mediates shoot Na⁺ exclusion, likely by withdrawing Na⁺ from the root xylem flow. A naturally occurring nonsynonymous SNP (SNP947-G) increases the Na⁺ transport activity of ZmHKT1;2, promoting shoot Na⁺ exclusion and salt tolerance in maize. SNP947-G first occurred in the wild grass teosinte (at a allele frequency of 43%) and has become a minor allele in the maize population (allele frequency 6.1%), suggesting that SNP947-G is derived from teosinte and that the genomic region flanking SNP947 likely has undergone selection during domestication or post-domestication dispersal of maize. Moreover, we demonstrate that introgression of the SNP947-G ZmHKT1;2 allele into elite maize germplasms reduces shoot Na⁺ content by up to 80% and promotes salt tolerance. Taken together, ZmNC3/ZmHKT1;2 was identified as an important QTL promoting shoot Na⁺ exclusion, and its favourable allele provides an effective tool for developing salt-tolerant maize varieties.