Arguments for the use of physiological criteria for improving the salt tolerance in crops

Efforts to develop new crop varieties with improved salt tolerance have been intensified over the past 15–20 years. Despite the existence of genetic variation for salt tolerance within species, and many methods available for expanding the source of genetic variation, there is only a limited number of varieties that have been developed with improved tolerance. These new varieties have all been based upon selection for agronomic characters such as yield or survival in saline conditions. That is, based upon characters that integrate the various physiological mechanisms responsible for tolerance. Yet over the same time period, knowledge of physiological salt responses has increased substantially.Selection and breeding to increase salt tolerance might be more successful if selection is based directly on the physiological mechanisms or characters conferring tolerance. Basic questions associated with using physiological selection criteria are discussed in the paper. These are centred around the need for genetic variation, the importance of the targeted mechanism, the ease of detection of the physiological mechanism (including the analytical requirements) and the breeding strategy. Many mechanisms, including ion exclusion, ion accumulation, compatible solute production and osmotic adjustment have been associated with genetic variation in salt tolerance. Yet their successful use in improving salt tolerance, via physiological selection criteria, is largely non-existent. Consideration is given to the role of physiological criteria in the short and long term in improving salt tolerance. In several glycophytic species, particularly legumes, physiological selection based on ion exclusion from the shoots shows promise. Recent results for white clover indicate the potential for using a broad physiological selection criterion of restricted Cl accumulation in the shoots, with scope for future refinement based upon the specific physiological characters that combined result in ion exclusion.     

Assessing the Suitability of Various Physiological Traits to Screen Wheat Genotypes for Salt Tolerance

Success of improving the salt tolerance of genotypes requires effective and reliable screening traits in breeding programs.The objective was to assess the suitability of various physiological traits to screen wheat genotypes for salt tolerance.Thirteen wheat genotypes from Egypt,Germany,Australia and India were grown in soil with two salinity levels(control and150mmol/L NaCl)in a greenhouse.The physiological traits(ion contents in leaves and stems,i.e.Na~ ,Cl~-,K~ ,Ca~(2 )),theratios of K~ /Na~ and Ca~ /Na~ in the leaves and stems,net photosynthesis rate,stomatal conductance,transpiration rate,chlorophyll content(SPAD value),and leaf water relations,were measured at different growth stages.The physiologicaltraits except for Na~ and Cl~-in stems and the leaf transpiration rate at 150 mmol/L NaCI showed a significant genotypicvariation,indicating that the traits that have a significant genotypic variation may be possibly used as screening criteria.According to the analysis of linear regression of the scores of the physiological traits against those of grain yield,however,the physiological traits of Ca~(2 )and Ca~(2 )/Na~ at 45 d and final harvest with the greatest genotypic variation were ranked atthe top.From a practical and economic point of view,SPAD value should be considered to be used as screening criteriaand/or there is a need to develop a quick and practical approach to determine Ca~(2 )in plant tissues.     

Assessing the Suitability of Various Physiological Traits to Screen Wheat Genotypes for Salt Tolerance

Success of improving the salt tolerance of genotypes requires effective and reliable screening traits in breeding programs. The objective was to assess the suitability of various physiological traits to screen wheat genotypes for salt tolerance. Thirteen wheat genotypes from Egypt, Germany, Australia and India were grown in soil with two salinity levels （control and 150 mmol/L NaCI） in a greenhouse. The physiological traits （ion contents in leaves and stems, i.e. Na^＋, Cl^-, K^＋, Ca^2＋）, the ratios of K^＋/Na^＋ and Ca^＋/Na^＋ in the leaves and stems, net photosynthesis rate, stomatal conductance, transpiration rate, chlorophyll content （SPAD value）, and leaf water relations, were measured at different growth stages. The physiological traits except for Na^＋ and Cl^- in stems and the leaf transpiration rate at 150 mmol/L NaCI showed a significant genotypic variation, indicating that the traits that have a significant genotypic variation may be possibly used as screening criteria. According to the analysis of linear regression of the scores of the physiological traits against those of grain yield, however, the physiological traits of Ca^2＋ and Ca2^＋/Na^＋ at 45 d and final harvest with the greatest genotypic variation were ranked at the top. From a practical and economic point of view, SPAD value should be considered to be used as screening criteria and/or there is a need to develop a quick and practical approach to determine Ca^2＋ in plant tissues.     

Halophyte Improvement for a Salinized World

It is more important to improve the salt tolerance of crops in a salinized world with the situations of increasing populations, declining crop yields, and a decrease in agricultural lands. Attempts to produce salt-tolerant crops have involved the manipulation of existing crops through conventional breeding, genetic engineering and marker-assisted selection (MAS). However, these have, so far, not produced lines growing on highly saline water. Hence, the domestication of wild halophytes as crops appears to be a feasible way to develop agriculture in highly saline environments. In this review, at first, the assessment criteria of salt tolerance for halophytes are discussed. The traditional criteria for the classification of salinity in crops are less applicable to strong halophytes with cubic growth curves at higher salinities. Thus, realistic assessment criteria for halophytes should be evaluated at low and high salinity levels. Moreover, absolute growth rather than relative growth in fields during a crop's life cycle should be considered. Secondly, the use of metabolomics to understand the mechanisms by which halophytes respond to salt tolerance is highlighted as is the potential for metabolomics-assisted breeding of this group of plants. Metabolomics provides a better understanding of the changes in cellular metabolism induced by salt stress. Identification of metabolic quantitative trait loci (QTL) associated with salt tolerance might provide a new method to aid the selection of halophyte improvement. Thirdly, the identification of germplasm-regression-combined (GRC) marker-trait association and its potential to identifying markers associated with salt tolerance is outlined. Results of MAS/linkage map-QTL have been modest because of the absence of QTLs with tight linkage, the non-availability of mapping populations and the substantial time needed to develop such populations. To overcome these limitations, identification by GRC-based marker-trait association has been successfully applied to many plant traits, including salt tolerance. Finally, we provide a prospect on the challenges and opportunities for halophyte improvement, especially in the integration of metabolomics- and GRC-marker-assisted selection towards new or unstudied halophyte breeding, for which no other genetic information, such as linkage maps and QTL, are available.     

马铃薯耐盐性的研究进展

土壤盐渍化已是影响全球农业生产和土地资源可持续利用的主要问题之一。近些年马铃薯的种植面积不断扩大,比重逐年上升,而马铃薯却是一种对盐中度敏感的作物,盐渍化对其造成严重的危害,因此研究马铃薯的耐盐性势在必行。本文对国内外马铃薯耐盐性筛选、生长发育、生理生化特性、耐盐途径、转基因耐盐马铃薯的研究及其二倍体马铃薯种质资源的利用和展望进行了综述。     

A review on transporters in salt tolerant mangroves

Key message

The role of transporters in imparting salt tolerance to mangroves is not yet understood. Identification of the role of transporters in halophytes is promising, as far as the development of genetically engineered salt tolerant crops is concerned.

Abstract

Mangroves are models for stress tolerance and they provide a reservoir for some of the novel genes and proteins, involved in salt tolerance. Biochemical or physiological mechanisms contribute to salt tolerance depending on variations in the environment. A great deal of research on salinity tolerance of plants, probes into water relations, photosynthesis, and accumulation of various in-organic ions and organic metabolites. The ability of the plant to react to high salinity depends on the genes that are expressed during stress. The mechanism of salinity tolerance becomes complicated when the responses of plants varies with salinity and environmental conditions. During the onset and development of salt stress within a plant, major processes such as photosynthesis, protein synthesis and lipid metabolisms are affected. The present review attempts to dissect out the role of transporters in salt tolerance of mangroves.     

Improving Salinity Tolerance in Cereals

Cereals are grown in almost every region of the world and are exposed to a variety of environmental stresses that severely affect their growth and grain yield. Of various abiotic stresses, salinity is one of the more significant threats to cereal crops. To ensure food security, there is a need to adopt strategies to overcome this specific threat. Undoubtedly, plant scientists have been exploiting a variety of approaches to achieve enhanced crop productivity on salt affected soils. Of the various biotic approaches, conventional breeding, marker-assisted selection and genetic engineering to develop salt-tolerant lines/cultivars of cereals all seem plausible. Some success stories have been reported for improvement in salt tolerance of wheat and rice, but are scarce for other cereals. A number of barriers to the development of salt-tolerant cultivars/lines have been identified and include a lack of knowledge about the genetics of crops, their physiological and biochemical behavior, wide variation in environmental conditions, and the complex polygenic nature of the salt tolerance character. This review focuses on how improvements have been made in salt tolerance in cereals through different biotic means, such as conventional breeding, marker assisted selection and genetic engineering.     

Hyperspectral imaging combined with machine learning as a tool to obtain high‐throughput plant salt‐stress phenotyping

The rapid selection of salinity‐tolerant crops to increase food production in salinized lands is important for sustainable agriculture. Recently, high‐throughput plant phenotyping technologies have been adopted that use plant morphological and physiological measurements in a non‐destructive manner to accelerate plant breeding processes. Here, a hyperspectral imaging (HSI) technique was implemented to monitor the plant phenotypes of 13 okra (Abelmoschus esculentus L.) genotypes after 2 and 7 days of salt treatment. Physiological and biochemical traits, such as fresh weight, SPAD, elemental contents and photosynthesis‐related parameters, which require laborious, time‐consuming measurements, were also investigated. Traditional laboratory‐based methods indicated the diverse performance levels of different okra genotypes in response to salinity stress. We introduced improved plant and leaf segmentation approaches to RGB images extracted from HSI imaging based on deep learning. The state‐of‐the‐art performance of the deep‐learning approach for segmentation resulted in an intersection over union score of 0.94 for plant segmentation and a symmetric best dice score of 85.4 for leaf segmentation. Moreover, deleterious effects of salinity affected the physiological and biochemical processes of okra, which resulted in substantial changes in the spectral information. Four sample predictions were constructed based on the spectral data, with correlation coefficients of 0.835, 0.704, 0.609 and 0.588 for SPAD, sodium concentration, photosynthetic rate and transpiration rate, respectively. The results confirmed the usefulness of high‐throughput phenotyping for studying plant salinity stress using a combination of HSI and deep‐learning approaches.     

Salinity Stress in Wheat: Effects,Mechanisms and Management Strategies

Salinity stress is a major threat to global food production and its intensity is continuously increasing because of anthropogenic activities. Wheat is a staple food and a source of carbohydrates and calories for the majority of people across the globe. However, wheat productivity is adversely affected by salt stress, which is associated with a reduction in germination, growth, altered reproductive behavior and enzymatic activity, disrupted photosynthesis, hormonal imbalance, oxidative stress, and yield reductions. Thus, a better understanding of wheat (plant) behavior to salinity stress has essential implications to devise counter and alleviation measures to cope with salt stress. Different approaches including the selection of suitable cultivars, conventional breeding, and molecular techniques can be used for facing salt stress tolerance. However, these techniques are tedious, costly, and labor-intensive. Management practices are still helpful to improve the wheat performance under salinity stress. Use of arbuscular mycorrhizal fungi, plant growth-promoting rhizobacteria, and exogenous application of phytohormones, seed priming, and nutrient management are important tools to improve wheat performance under salinity stress. In this paper, we discussed the effect of salinity stress on the wheat crop, possible mechanisms to deal with salinity stress, and management options to improve wheat performance under salinity conditions.     

Salinity Tolerance in Brassica Oilseeds

Brassica oilseed species now hold the third position among oilseed crops and are an important source of vegetable oil. The most common Brassica oil-seed crops grown for commercial purposes are rape seeds, (Brassica campestris L. and B. napus L.) and mustards (B. juncea (L.) Czern. & Coss. and B. carinata A.Br.). The other Brassica species such as B. nigra (L.) Koch and B. tournefortii Gouan are grown on a very small scale. Brassica napus, B. juncea, and B. carinata are amphidiploids, whereas B. campestris and B. nigra are diploid. Most of the Brassica species have been categorized as moderately salt tolerant, with the amphidiploid species being the relatively salt tolerant in comparison with the diploid species. Due to the higher salt tolerance of the amphidiploids, it has been suggested that their salt tolerance has been acquired from the A (B. campestris) and C (B. oleracea L.) genomes. However, significant inter- and intraspecific variation for salt tolerance exists within brassicas, which can be exploited through selection and breeding for enhancing salt tolerance of the crops. There are contrasting reports regarding the response of these species to salinity at different plant developmental stages, but in most of them it is evident that they maintain their degree of salt tolerance consistently throughout the plant ontogeny. The pattern of uptake and accumulation of toxic ions (Na⁺ and Cl^?), in tissues of plants subjected to saline conditions appears to be mostly due to mechanism of partial ion exclusion (exclusion of Na⁺ and/or Cl^?) in most of the species, although ion inclusion in some cases at intraspecific levels has also been observed. Maintenance of high tissue K⁺/Na⁺ and Ca^{2 +}/Na⁺ ratios has been suggested as an important selection criterion for salt-tolerance in brassicas. Osmotic adjustment has also been reported in Brassica plants subjected to saline conditions, but particularly to a large extent in salt-tolerant species or cultivars. The roles of important organic osmotica such as total soluble sugars, free amino acids, and free proline, which are central to osmotic adjustment, have been discussed. In canola, B. napus, no positive relationship has been observed between salt tolerance and erucic acid content of seed oil in different cultivars. Furthermore, glucosinolate content of the seed meal in canola generally increases with an increase in salt level of the growth medium. This review highlights the responses of potential Brassica crops to soil salinity from the whole plant to the molecular level. It also describes the efforts made during the past millennium in uncovering the mechanism(s) of salinity tolerance of these crops both at the whole plant and cellular levels. The important selection criteria, which are used by researchers to enhance the degree of salinity tolerance in brassicas, are summarized. In addition, the vital role of genetic engineering and molecular biology approaches to the improvement of salt tolerance in brassicas is emphasized.     

Comparative roles of brassinosteroids and polyamines in salt stress tolerance

Salt stress is among the major abiotic stresses that adversely affect the global crop production and its adverse impacts are getting more serious in the regions where saline water is used for irrigation. It induces reactive oxygen species, alters the activity of antioxidant system and adversely affects the process of photosynthesis. Various strategies have been employed to mitigate the deleterious effects of salt stress. Presently, the recommended strategies to overcome the adverse effects of salt stress include the use of tolerant cultivars, ameliorative water management and diverse cultural practices. However, none of these approaches have been found to be fully effective under salt stress conditions. An alternative and technically simpler approach to induce salt stress tolerance is the exogenous application of plant growth regulators (PGRs). This technique has gained significant importance during the past decade. PGRs have been implicated to regulate a wide range of metabolic and physiological activities in plants, ranging from cell division and organogenesis to protection against biotic and abiotic stresses. One of the important factors for enhanced plant productivity by PGRs is their efficiency to overcome the salt-induced stress conditions. Recent findings on the effects of brassinosteroids and polyamines on the salt stress tolerance of crops open new avenues to address the salinity problems. This review enlightens the role of brassinosteroids and polyamines in different plant processes like their role in regulation of photosynthesis, antioxidant systems and other related aspects, thereby improving overall performance of plants.     

Salt-tolerant crops: origins,development, and prospects of the concept

Summary The genetic approach to the problems posed by salt-affected soils and water,i.e., breeding crops resistant to salinity stress, is traced to two principal origins: the European ecological interest in halophytes, and the exigencies of growing crops in the arid and semi-arid lands of the American West. The point is made that breeding for resistance to salinity stress cannot be divorced from breeding for various other desirable traits of mineral plant nutrition and metabolism. A survey is conducted of the existing body of information on breeding for desiderata of mineral nutrition in general and salt tolerance in particular. The prospects of breeding crops for salt tolerance are discussed, with emphasis on a) its relation to breeding for resistance to other mineral stresses; b) field trials; c) collaboration between plant physiologists and geneticist-breeders; and d) extensive exploration of germplasm.     

野生大豆耐盐性研究进展

野生大豆对于拓宽大豆种质遗传基础和丰富大豆种质基因库具有重要意义.该文从野生大豆的资源概况及优良性状、耐盐机理和利用野生大豆提高栽培大豆耐盐性等三个方面,对近年来国内外有关野生大豆耐盐性的解剖结构、生理基础、分子生物学基础等方面的研究进展进行了系统综述,并提出野生大豆通过茎叶表皮上的"腺体"及对Na+和Cl-的排斥性,实现对盐渍环境的颉颃作用.较强的抗氧化能力、大豆异黄酮代谢和耐盐基因也是其适应盐渍环境的重要原因.今后应对野生大豆耐盐机理的遗传学基础进行深入研究,并通过种群保护以保障野生大豆的发掘鉴定和创新利用.     

Breeding for abiotic stresses for sustainable agriculture

Using cereal crops as examples, we review the breeding for tolerance to the abiotic stresses of low nitrogen, drought, salinity and aluminium toxicity. All are already important abiotic stress factors that cause large and widespread yield reductions. Drought will increase in importance with climate change, the area of irrigated land that is salinized continues to increase, and the cost of inorganic N is set to rise. There is good potential for directly breeding for adaptation to low N while retaining an ability to respond to high N conditions. Breeding for drought and salinity tolerance have proven to be difficult, and the complex mechanisms of tolerance are reviewed. Marker-assisted selection for component traits of drought in rice and pearl millet and salinity tolerance in wheat has produced some positive results and the pyramiding of stable quantitative trait locuses controlling component traits may provide a solution. New genomic technologies promise to make progress for breeding tolerance to these two stresses through a more fundamental understanding of underlying processes and identification of the genes responsible. In wheat, there is a great potential of breeding genetic resistance for salinity and aluminium tolerance through the contributions of wild relatives.     

Improving salinity tolerance in crop plants: a biotechnological view

Salinity limits the production capabilities of agricultural soils in large areas of the world. Both breeding and screening germplasm for salt tolerance encounter the following limitations: (a) different phenotypic responses of plants at different growth stages, (b) different physiological mechanisms, (c) complicated genotype × environment interactions, and (d) variability of the salt-affected field in its chemical and physical soil composition. Plant molecular and physiological traits provide the bases for efficient germplasm screening procedures through traditional breeding, molecular breeding, and transgenic approaches. However, the quantitative nature of salinity stress tolerance and the problems associated with developing appropriate and replicable testing environments make it difficult to distinguish salt-tolerant lines from sensitive lines. In order to develop more efficient screening procedures for germplasm evaluation and improvement of salt tolerance, implementation of a rapid and reliable screening procedure is essential. Field selection for salinity tolerance is a laborious task; therefore, plant breeders are seeking reliable ways to assess the salt tolerance of plant germplasm. Salt tolerance in several plant species may operate at the cellular level, and glycophytes are believed to have special cellular mechanisms for salt tolerance. Ion exclusion, ion sequestration, osmotic adjustment, macromolecule protection, and membrane transport system adaptation to saline environments are important strategies that may confer salt tolerance to plants. Cell and tissue culture techniques have been used to obtain salt tolerant plants employing two in vitro culture approaches. The first approach is selection of mutant cell lines from cultured cells and plant regeneration from such cells (somaclones). In vitro screening of plant germplasm for salt tolerance is the second approach, and a successful employment of this method in durum wheat is presented here. Doubled haploid lines derived from pollen culture of F₁ hybrids of salt-tolerant parents are promising tools to further improve salt tolerance of plant cultivars. Enhancement of resistance against both hyper-osmotic stress and ion toxicity may also be achieved via molecular breeding of salt-tolerant plants using either molecular markers or genetic engineering.     

Research Advances in Mechanisms of Turfgrass Tolerance to Abiotic Stresses: From Physiology to Molecular Biology

Turgrfass used on landscapes, parks, sports fields, and golf courses has significant ecological, environmental, and economic impacts. The economic value of seed production of turfgrasses is second to hybrid corn. The land area cultivated with turfgrass is increasing due to rapid urban development. Turfgrass is often subjected to various abiotic stresses, which cause declines in aesthetic quality, functionality and seed yield. Among abiotic stresses, drought, salinity, heat, and low temperature are the most common detrimental factors for turfgrass growth in various regions. Thorough understanding of mechanisms of turfgrass stress responses is vital for the development of superior stress-tolerant germplasm through breeding and biotechnology. Significant progress has been made in turfgrass stress physiology and molecular biology in recent decades, but research for turfgrasses generally lags behind that of the major Poaceae crops, particularly at the molecular and genomic levels. This review focuses on research advances in turfgrass stress physiology and provides an overview of limited information on gene discovery, genetic transformation, and molecular marker development for improving stress tolerance, with emphasis on drought, salinity, heat, and low temperature stress. Major growth and physiological traits associated with these stresses, as well as metabolic and molecular factors regulating various traits for turfgrass tolerance to each stress are discussed. Future research at the systems biology level and through genomic sequencing is paramount for further insights on fundamental mechanisms of turfgrass stress tolerance and for improving turfgrass tolerance to various environmental stresses.     

Principles and strategies in breeding for higher salt tolerance

Summary Salinity is an environmental component that usually reduces yield. Recent advances in the understanding of salt effects on plants have not revealed a reliable physiological or biochemical marker that can be used to rapidly screen for salt tolerance. The necessity of measuring salt tolerance based upon growth in saline relative to non-saline environments makes salt tolerance measurements and selection for tolerance difficult. Additionally, high variability in soil salinity and environmental interactions makes it questionable whether breeding should be conducted for tolerance or for high yield. Genetic techniques can be used to identify the components of variation attributable to genotype and environment, and the extent of genetic variation in saline and nonsaline environments can be used to estimate the potential for improving salt tolerance. Absolute salt tolerance can be improved best by increasing both absolute yield and relative salt tolerance.     

Avenues for increasing salt tolerance of crops,and the role of physiologically based selection traits

Increased salt tolerance is needed for crops grown in areas at risk of salinisation. This requires new genetic sources of salt tolerance, and more efficient techniques for identifying salt-tolerant germplasm, so that new genes for tolerance can be introduced into crop cultivars. Screening a large number of genotypes for salt tolerance is not easy. Salt tolerance is achieved through the control of salt movement into and through the plant, and salt-specific effects on growth are seen only after long periods of time. Early effects on growth and metabolism are likely due to osmotic effects of the salt, that is to the salt in the soil solution. To avoid the necessity of growing plants for long periods of time to measure biomass or yield, practical selection techniques can be based on physiological traits. We illustrate this with current work on durum wheat, on selection for the trait of sodium exclusion. We have explored a wide range of genetic diversity, identified a new source of sodium exclusion, confirmed that the trait has a high heritability, checked for possible penalties associated with the trait, and are currently developing molecular markers. This illustrates the potential for marker-assisted selection based on sound physiological principles in producing salt-tolerant crop cultivars.     

Modeling uptake of cadmium from solution outside of root to cell wall of shoot in rice seedling

Barley (Hordeum vulgare L.) is well known for its relatively high salt tolerance among cereal crops. However, the genetic variation of cultivated barley becomes narrower due to continuous artificial selection and breeding processes. Compared with cultivated barley, wild barley contains wider genetic variation and abundant sources for abiotic stress tolerance, considering as an elite resource for mechanism study on salt tolerance. In this study, Tibetan wild barley accession XZ113 identified with high salt tolerance, was used to investigate ionic responses and to identify proteins involved in salt tolerance in roots and shoots at early stage of salt stress, during 48 h. Exposed to salinity, shoot growth is more sensitive than root growth. Conversely, K/Na ratio in the shoots was larger than that in the roots, and both were above 1.0. Steady-state K⁺ flux experiment showed XZ113 had a strong K⁺-retaining ability under salt stress, maybe contributing to its good performance of the absolute growth rate. Proteomic results suggested that monodehydroascorbate reductase and peroxidases related to reactive oxygen species scavenging in the roots and phosphoglycerate kinase, triosephosphate isomerase and sedoheptulose-1,7-bisphosphatase associated with photosynthesis and metabolisms in the shoots, played important roles in salt tolerance at early stage of salinity in wild barley.