Induction of abiotic stress tolerance in plants by endophytic microbes

Endophytes are micro‐organisms including bacteria and fungi that survive within healthy plant tissues and promote plant growth under stress. This review focuses on the potential of endophytic microbes that induce abiotic stress tolerance in plants. How endophytes promote plant growth under stressful conditions, like drought and heat, high salinity and poor nutrient availability will be discussed. The molecular mechanisms for increasing stress tolerance in plants by endophytes include induction of plant stress genes as well as biomolecules like reactive oxygen species scavengers. This review may help in the development of biotechnological applications of endophytic microbes in plant growth promotion and crop improvement under abiotic stress conditions.

Significance and Impact of the Study

Increasing human populations demand more crop yield for food security while crop production is adversely affected by abiotic stresses like drought, salinity and high temperature. Development of stress tolerance in plants is a strategy to cope with the negative effects of adverse environmental conditions. Endophytes are well recognized for plant growth promotion and production of natural compounds. The property of endophytes to induce stress tolerance in plants can be applied to increase crop yields. With this review, we intend to promote application of endophytes in biotechnology and genetic engineering for the development of stress‐tolerant plants.     

Role of microorganisms in adaptation of agriculture crops to abiotic stresses

Increased incidences of abiotic and biotic stresses impacting productivity in principal crops are being witnessed all over the world. Extreme events like prolonged droughts, intense rains and flooding, heat waves and frost damages are likely to further increase in future due to climate change. A wide range of adaptations and mitigation strategies are required to cope with such impacts. Efficient resource management and crop/livestock improvement for evolving better breeds can help to overcome abiotic stresses to some extent. However, such strategies being long drawn and cost intensive, there is a need to develop simple and low cost biological methods for the management of abiotic stress, which can be used on short term basis. Microorganisms could play a significant role in this respect, if we can exploit their unique properties of tolerance to extremities, their ubiquity, genetic diversity, their interaction with crop plants and develop methods for their successful deployment in agriculture production. Besides influencing the physico-chemical properties of rhizospheric soil through production of exopolysaccharides and formation of biofilm, microorganisms can also influence higher plants response to abiotic stresses like drought, chilling injury, salinity, metal toxicity and high temperature, through different mechanisms like induction of osmo-protectants and heat shock proteins etc. in plant cells. Use of these microorganisms per se can alleviate stresses in crop plants thus opening a new and emerging application in agriculture. These microbes also provide excellent models for understanding the stress tolerance, adaptation and response mechanisms that can be subsequently engineered into crop plants to cope with climate change induced stresses.     

Physiological adaptive mechanisms of plants grown in saline soil and implications for sustainable saline agriculture in coastal zone

There is large area of saline abandoned and low-yielding land distributed in coastal zone in the world. Soil salinity which inhibits plant growth and decreases crop yield is a serious and chronic problem for agricultural production. Improving plant salt tolerance is a feasible way to solve this problem. Plant physiological and biochemical responses under salinity stress become a hot issue at present, because it can provide insights into how plants may be modified to become more tolerant. It is generally known that the negative effects of soil salinity on plants are ascribed to ion toxicity, oxidative stress and osmotic stress, and great progress has been made in the study on molecular and physiological mechanisms of plant salinity tolerance in recent years. However, the present knowledge is not easily applied in the agronomy research under field environment. In this review, we simplified the physiological adaptive mechanisms in plants grown in saline soil and put forward a practical procedure for discerning physiological status and responses. In our opinion, this procedure consists of two steps. First, negative effects of salt stress are evaluated by the changes in biomass, crop yield and photosynthesis. Second, the underlying reasons are analyzed from osmotic regulation, antioxidant response and ion homeostasis. Photosynthesis is a good indicator of the harmful effects of saline soil on plants because of its close relation with crop yield and high sensitivity to environmental stress. Particularly, chlorophyll a fluorescence transient has been accepted as a reliable, sensitive and convenient tool in photosynthesis research in recent years, and it can facilitate and enrich photosynthetic research under field environment.     

Drought-induced responses of photosynthesis and antioxidant metabolism in higher plants

Environmental stresses trigger a wide variety of plant responses, ranging from altered gene expression and cellular metabolism to changes in growth rates and crop yields. A plethora of plant reactions exist to circumvent the potentially harmful effects caused by a wide range of both abiotic and biotic stresses, including light, drought, salinity, high temperatures, and pathogen infections. Among the environmental stresses, drought stress is one of the most adverse factors of plant growth and productivity. Understanding the biochemical and molecular responses to drought is essential for a holistic perception of plant resistance mechanisms to water-limited conditions. Drought stress progressively decreases CO2 assimilation rates due to reduced stomatal conductance. Drought stress also induces reduction in the contents and activities of photosynthetic carbon reduction cycle enzymes, including the key enzyme, ribulose-1,5-bisphosphate carboxylase/oxygenase. The critical roles of proline and glycine-betaine, as well as the role of abscisic acid (ABA), under drought stress conditions have been actively researched to understand the tolerance of plants to dehydration. In addition, drought stress-induced generation of active oxygen species is well recognized at the cellular level and is tightly controlled at both the production and consumption levels in vivo, through increased antioxidative systems. Knowledge of sensing and signaling pathways, including ABA-mediated changes in response to drought stress, is essential to improve crop management. This review focuses on the ability and strategies of higher plants to respond and adapt to drought stress.     

Role of Triacontanol in Counteracting the Ill Effects of Salinity in Plants: A Review

Soil salinity is one among the common environmental threats to agriculture. It adversely affects the physio-biochemical processes of plants that eventually lead to the reduction in growth, development and crop productivity. To cope with such adverse conditions, plants develop certain internal mechanisms, but under severe conditions these mechanisms fail to tolerate the salt stress. To overcome this problem, various strategies have been employed that help plants to mitigate salinity effects. Among the various strategies, the application of plant growth regulators (PGRs) has gained significant attention to induce salt tolerance in plants. A number of PGRs have been used so far. Among these, triacontanol (TRIA), a new PGR is gaining a lot of importance to enhance the plant growth, productivity and salinity tolerance in different crops. The utility of TRIA is dependent on its applied concentration. Its lower concentrations generally alleviate the salinity effects. However, the knowledge of its biosynthesis, signalling and its role particularly to mitigate salinity effect remains scanty. In the present article, the focus has been given on the role of exogenous applications of TRIA in the regulation of physio-biochemical characteristics especially plant growth, photosynthesis, nutrient acquisition, oxidative stress, antioxidant systems, compatible solutes, yield attributes and its mode of action in plants under salinity conditions. The salient features of the review may provide new insights on the role of TRIA in countering the ill effect of salinity in different crop plants.

     

Cadmium stress tolerance in crop plants: Probing the role of sulfur

Plants can''t move away and are therefore continuously confronted with unfavorable environmental conditions (such as soil salinity, drought, heat, cold, flooding and heavy metal contamination). Among heavy metals, cadmium (Cd) is a non-essential and toxic metal, rapidly taken up by roots and accumulated in various plant tissues which hamper the crop growth and productivity worldwide. Plants employ various strategies to counteract the inhibitory effect of Cd, among which nutrient management is one of a possible way to overcome Cd toxicity. Sulfur (S) uptake and assimilation are crucial for determining crop yield and resistance to Cd stress. Cd affects S assimilation pathway which leads to the activation of pathway responsible for the synthesis of cysteine (Cys), a precursor of glutathione (GSH) biosynthesis. GSH, a non-protein thiol acts as an important antioxidant in mitigating Cd-induced oxidative stress. It also plays an important role in phytochelatins (PCs) synthesis, which has a proven role in Cd detoxification. Therefore, S assimilation is considered a crucial step for plant survival under Cd stress. The aim of this review is to discuss the regulatory mechanism of S uptake and assimilation, GSH and PC synthesis for Cd stress tolerance in crop plants.Key words: cadmium, cysteine, glutathione, phytochelatins, stress tolerance, sulfur     

Crop production and management under saline conditions

Summary This review evaluates management practices that may minimize yield reduction under saline conditions according to three strategies: (I) control of root-zone salinity; (II) reduced damage to the crop; (III) reduced damage to individual plants. Plant response to salinity is described by an unchanged yield up to a threshold soil salinity (a), then a linear reduction in relative yield (b), to a maximum soil salinity that corresponds to zero yield (Y_o). Strategies I and II do not take into consideration any change in the parameters of the response curve, while strategy III is aimed at modifying them.Control of root zone salinity is obtained by irrigation and leaching. From the review of existing data it is concluded that the effective soil salinity parameter should be taken as the mean electrical conductivity of the saturated paste extract or of the soil solution over time and space. Several irrigation and leaching practices are discussed. It is shown that intermittent leaching is more advantageous than leaching at each irrigation. Specific cultivation and irrigation practices that result in soil salinity reduction adjacent to young seedlings and the use of water of low salinity at specifically sensitive growth stages may be highly beneficial. Recent data do not show that reduced irrigation intervals improve crop response more under saline than under nonsaline irrigation. Alternate use of water of different salt concentrations results in mixing in the soil and the crop responds to the mean water salinity.Reduced damage at the fiel level when soil or irrigation water salinity is too high to maintain full yield of single plants requires a larger crop stand. For row crops reduced inter-row spacing is more effective than reduced intra-row spacing.Reduced damage at the plant level while the salinity tolerance of the plants remains constant shows up in the response curve parameters as larger threshold and slope and constant salinity at zero yield. This is the effect of a reduced atmospheric water demand that results in reduced stress in the plant under given salinity. Management can also change the salt tolerance of the crop. This will show up as higher salinity at zero yield, as well as changes in threshold and slope. Such changes in the response curve were found at different growth stages, under different atmospheric CO₂, under different fertilization, and when sprinkler irrigation was compared with drip irrigation.Contribution from the Agricultural Research Organization, The Volcani Center, Bet Dagan, Israel. No. 1111-E 1984 series.     

Connecting salt stress signalling pathways with salinity‐induced changes in mitochondrial metabolic processes in C3 plants

Salinity exerts a severe detrimental effect on crop yields globally. Growth of plants in saline soils results in physiological stress, which disrupts the essential biochemical processes of respiration, photosynthesis, and transpiration. Understanding the molecular responses of plants exposed to salinity stress can inform future strategies to reduce agricultural losses due to salinity; however, it is imperative that signalling and functional response processes are connected to tailor these strategies. Previous research has revealed the important role that plant mitochondria play in the salinity response of plants. Review of this literature shows that 2 biochemical processes required for respiratory function are affected under salinity stress: the tricarboxylic acid cycle and the transport of metabolites across the inner mitochondrial membrane. However, the mechanisms by which components of these processes are affected or react to salinity stress are still far from understood. Here, we examine recent findings on the signal transduction pathways that lead to adaptive responses of plants to salinity and discuss how they can be involved in and be affected by modulation of the machinery of energy metabolism with attention to the role of the tricarboxylic acid cycle enzymes and mitochondrial membrane transporters in this process.     

Comprehensive insight into arbuscular mycorrhizal fungi,Trichoderma spp. and plant multilevel interactions with emphasis on biostimulation of horticultural crops

Sustainability and a more environment-friendly approach is an emerging issue relevant to crop production. Abiotic stresses like drought, salinity, heat, cold or heavy metal pollution can severely compromise yields, and in this respect, plant protection practices should be highly efficient as well as safe for the environment and people. Among the many ways to achieve high productivity of healthy, safe and tasty food, the use of beneficial micro-organisms as biostimulants is the most promising one. Two types of soil fungi can be considered efficient natural plants stimulants: arbuscular mycorrhizal fungi (AMF) and Trichoderma spp. (TR). Generally, most investigations indicated AMF and TR were effective, as well as safe, for use as natural biopreparations dedicated to horticultural crops, although some reports pointed to their negative impact on plants. This review focuses on the mutual interaction of AMF and TR, as well as complex relationships with plants analysed on a multidimensional level: biochemical, morphological, ecological and agrotechnical. AMF and TR were found to be effective elicitors of root system development, nutrient uptake, plant stress response and production of secondary metabolites. As natural plant stimulants, beneficial fungi are compatible with modern trends of crop management, environmental conservation and functional food production. Herein, we demonstrate the advantages and disadvantages of AMF and TR use in horticulture and their prospects, as well as the points that need further exploring.     

Soil and crop management strategies to prevent iron deficiency in crops

Plants and humans cannot easily acquire iron from their nutrient sources although it is abundant in nature. Thus, iron deficiency is one of the major limiting factors affecting crop yields, food quality and human nutrition. Therefore, approaches need to be developed to increase Fe uptake by roots, transfer to edible plant portions and absorption by humans from plant food sources. Integrated strategies for soil and crop management are attractive not only for improving growing conditions for crops but also for exploiting a plant??s potential for Fe mobilization and utilization. Recent research progress in soil and crop management has provided the means to resolve complex plant Fe nutritional problems through manipulating the rhizosphere (e.g., rhizosphere fertilization and water regulation), and crop management (includes managing cropping systems and screening for Fe efficient species and varieties). Some simple and effective soil management practices, termed ??rhizosphere fertilization?? (such as root feeding and bag fertilization) have been developed and widely used by local farmers in China to improve the Fe nutrition of fruit plants. Production practices for rice cultivation are shifting from paddy-rice to aerobic rice to make more efficient use of irrigation water. This shift has brought about increases in Fe deficiency in rice, a new challenge depressing iron availability in rice and reducing Fe supplies to humans. Current crop management strategies addressing Fe deficiency include Fe foliar application, trunk injection, plant breeding for enriched Fe crop species and varieties, and selection of cropping systems. Managing cropping systems, such as intercropping strategies may have numerous advantages in terms of increasing Fe availability to plants. Studies of intercropping systems on peanut/maize, wheat/chickpea and guava/sorghum or -maize increased Fe content of crops and their seed, which suggests that a reasonable intercropping system of iron-efficient species could prevent or mitigate Fe deficiency in Fe-inefficient plants. This review provides a comprehensive comparison of the strategies that have been developed to address Fe deficiency and discusses the most recent advance in soil and crop management to improve the Fe nutrition of crops. These proofs of concept studies will serve as the basis for future Fe research and for integrated and optimized management strategies to alleviate Fe deficiency in farmers?? fields.     

Increasing the okra salt threshold value with biochar amendments

Soil salinity is a severe worldwide environmental problem that adversely affects soil properties and the crop growth such as okra. We hypothesized that biochar soil amendments could increase the okra salt threshold, alleviate salt stress and improve soil productivity. In this study, a pot experiment was conducted to investigate whether biochar could ameliorate the effects of salinity on okra plants. Three biochar amendment (BA) soil applications (0%, 5% and 10% by mass of soil) were considered for seven irrigation water salinity levels (0.75, 1.0, 2.0, 4.0, 5.0, 6.0 and 7.0?dS?m^?1) in a randomized block design with three replications. The Maas and Hoffman salt tolerance model was used to evaluate the effects of BA on okra plant growth parameters (e.g. yield, biomass) and water use efficiency for each salinity treatment. The results showed that increasing the soil salinity levels caused significant decreases in plant yields and yield components. However, biochar application rates of 5% and 10% increased the okra threshold by 19.7% and 81.2%, respectively, compared to the control (0%). The 10% biochar application rate also resulted in the greatest okra plant growth and increased yield, indicating that the effects of salt stress were ameliorated; moreover, the soil bulk density was decreased, and the water content was increased. Hence, biochar soil amendments could be considered as an important agronomic practice that could potentially overcome the adverse effects of salt stress.     

Halophytes as a source of genes for abiotic stress tolerance

Agricultural productivity is majorly impacted due to various abiotic stresses, particularly salinity and drought. Halophytes serve as an excellent resource for identifying and developing new crop systems, as these grow very luxuriously in very high saline soils. Understanding salinity stress tolerance mechanisms in such plants is an important step towards generating crop varieties that can cope with environmental stresses. Use of modern tools of ‘omics’ analyses and small RNA sequencing has helped to gain insights into the complex plant stress responses. Salinity tolerance being a multigenic trait requires a combination of strategies and techniques to successfully develop improved crops varieties. Many transgenic crops are being developed through genetic transformation. Besides marker-assisted breeding/QTL approaches are also being used to improve abiotic stress tolerance. In this review, we focus on the recent developments in the utilization of halophytes as a source of genes for genetic improvement in abiotic stress tolerance of crops.     

Salt stress tolerance; what do we learn from halophytes?

Abiotic environmental stresses can give rise to morphological, biochemical and molecular changes that negatively affect plant growth and productivity. Among these stresses, soil salinity is the major threat. To deal and control effects of high salinity on plants, it is important to understand their responses to salt stress that disturbs the homeostatic equilibrium at cellular and molecular levels. In this regard halophytes (salt tolerant plants) can provide superior models for the study of salt stress defense parameters compared to salt sensitive species (glycophytes). Halophytes use highly developed, complex systems to tolerate salinity by maintaining a low cytosolic Na⁺/K⁺ ratio, sequestration of Na⁺ into vacuoles that then provides the osmotic potential sustaining water influx. Under low intensity stress conditions that moderately and/or transiently affect ion imbalance, the set of responses all plants initiate will be mostly to engage measures that assure ion balance. High salinity, especially over a prolonged time period, will challenge plant survival, which then requires different strategies that employ a variety of mechanisms. Plasticity and connectivity of these diverse mechanisms is engrained in species- and family-specific evolutionary history and their genetic complexity. Highlighting differences in the genetic and biochemical makeup between glycophytes and halophytes allows for comparisons between their approaches towards high salinity. This review provides a brief overview about different strategies and mechanism used by plants to avoid or confine adverse effects of high salinity.     

Use of plant growth-promoting rhizobacteria to control stress responses of plant roots

Ethylene is a key gaseous hormone that controls various physiological processes in plants including growth, senescence, fruit ripening, and responses to abiotic and biotic stresses. In spite of some of these positive effects, the gas usually inhibits plant growth. While chemical fertilizers help plants grow better by providing soil-limited nutrients such as nitrogen and phosphate, over-usage often results in growth inhibition by soil contamination and subsequent stress responses in plants. Therefore, controlling ethylene production in plants becomes one of the attractive challenges to increase crop yields. Some soil bacteria among plant growth-promoting rhizobacteria (PGPRs) can stimulate plant growth even under stressful conditions by reducing ethylene levels in plants, hence the term “stress controllers” for these bacteria. Thus, manipulation of relevant genes or gene products might not only help clear polluted soil of contaminants but contribute to elevating the crop productivity. In this article, the beneficial soil bacteria and the mechanisms of reduced ethylene production in plants by stress controllers are discussed.     

Endophytic Bacteria in Plant Salt Stress Tolerance: Current and Future Prospects

Soil salinity is a major limiting factor for crop productivity worldwide and is continuously increasing owing to climate change. A wide range of studies and practices have been performed to induce salt tolerance mechanisms in plants, but their result in crop improvement has been limited due to lack of time and money. In the current scenario, there is increasing attention towards habitat-imposed plant stress tolerance driven by plant-associated microbes, either rhizospheric and/or endophytic. These microbes play a key role in protecting plants against various environmental stresses. Therefore, the use of plant growth-promoting microbes in agriculture is a low-cost and eco-friendly technology to enhance crop productivity in saline areas. In the present review, the authors describe the functionality of endophytic bacteria and their modes of action to enhance salinity tolerance in plants, with special reference to osmotic and ionic stress management. There is concrete evidence that endophytic bacteria serve host functions, such as improving osmolytes, anti-oxidant and phytohormonal signaling and enhancing plant nutrient uptake efficiency. More research on endophytes has enabled us to gain insights into the mechanism of colonization and their interactions with plants. With this information in mind, the authors tried to solve the following questions: (1) how do benign endophytes ameliorate salt stress in plants? (2) What type of physiological changes incur in plants under salt stress conditions? And (3), what type of determinants produced by endophytes will be helpful in plant growth promotion under salt stress?

     

荒漠植物不同功能群性状特征及其与土壤环境的关系

研究不同功能群植物性状差异及其与土壤环境关系对于充分掌握植物的环境适应策略至关重要。以艾比湖流域为研究区,利用荒漠植物的植物高度、叶片碳、氮、磷、硫、钾、钙、钠、镁含量等9个性状,将高、低土壤水盐环境下的植物划分为5个功能群,分析不同功能群的植物组成、性状差异及其与土壤环境的关系。结果表明：(1)不同土壤水盐环境下,其植物功能群组成不同;其中白刺、胡杨和罗布麻在两个土壤水盐环境下的功能群中均存在。(2)植物的功能性状在不同土壤水盐环境下也发生了适应性的变化。高土壤水盐环境下3个功能群的植物高度、叶片碳、氮、磷和钙含量显著高于低土壤水盐环境功能群(P<0.05);低土壤水盐环境下2个功能群的植物叶片硫、钠和镁含量高于高土壤水盐环境功能群。(3)土壤含水量(SVWC)、电导率(EC)、pH以及土壤磷含量对荒漠植物功能性状影响较大。在高土壤水盐环境下,EC、pH与植物高度,叶片钾、钙含量正相关,与叶片硫含量负相关;在低土壤水盐环境下,SVWC、EC与植物高度呈显著正相关(P<0.05)。研究有助于理解荒漠植物对极端环境的适应对策,为保护荒漠地区生物多样性提供理论依据。     

The pepper late embryogenesis abundant protein CaLEA1 acts in regulating abscisic acid signaling,drought and salt stress response

As sessile organisms, plants are constantly challenged by environmental stresses, including drought and high salinity. Among the various abiotic stresses, osmotic stress is one of the most important factors for growth and significantly reduces crop productivity in agriculture. Here, we report a function of the CaLEA1 protein in the defense responses of plants to osmotic stress. Our analyses showed that the CaLEA1 gene was strongly induced in pepper leaves exposed to drought and increased salinity. Furthermore, we determined that the CaLEA1 protein has a late embryogenesis abundant (LEA)_3 homolog domain highly conserved among other known group 5 LEA proteins and is localized in the processing body. We generated CaLEA1‐silenced peppers and CaLEA1‐overexpressing (OX) transgenic Arabidopsis plants to evaluate their responses to dehydration and high salinity. Virus‐induced gene silencing of CaLEA1 in pepper plants conferred enhanced sensitivity to drought and salt stresses, which was accompanied by high levels of lipid peroxidation in dehydrated and NaCl‐treated leaves. CaLEA1‐OX plants exhibited enhanced sensitivity to abscisic acid (ABA) during seed germination and in the seedling stage; furthermore, these plants were more tolerant to drought and salt stress than the wild‐type plants because of enhanced stomatal closure and increased expression of stress‐responsive genes. Collectively, our data suggest that CaLEA1 positively regulates drought and salinity tolerance through ABA‐mediated cell signaling.     

Silicon: a duo synergy for regulating crop growth and hormonal signaling under abiotic stress conditions

Abiotic stresses, such as salinity, heavy metals and drought, are some of the most devastating factors hindering sustainable crop production today. Plants use their own defensive strategies to cope with the adverse effects of these stresses, via the regulation of the expression of essential phytohormones, such as gibberellins (GA), salicylic acid (SA), jasmonates (JA), abscisic acid (ABA) and ethylene (ET). However, the efficacy of the endogenous defensive arsenals of plants often falls short if the stress persists over an extended period. Various strategies are developed to improve stress tolerance in plants. For example, silicon (Si) is widely considered to possess significant potential as a substance which ameliorate the negative effects of abiotic stresses, and improves plant growth and biomass accumulation. This review aims to explain how Si application influences the signaling of the endogenous hormones GA, SA, ABA, JA and ET during salinity, wounding, drought and metal stresses in crop plants. Phytohormonal cross talk plays an important role in the regulation of induced defences against stress. However, detailed molecular and proteomic research into these interactions is needed in order to identify the underlying mechanisms of stress tolerance that is imparted by Si application and uptake.