Targeting Glycinebetaine for Abiotic Stress Tolerance in Crop Plants: Physiological Mechanism,Molecular Interaction and Signaling

In the era of climate change, abiotic stresses (e.g., salinity, drought, extreme temperature, flooding, metal/metalloid(s), UV radiation, ozone, etc.) are considered as one of the most complex environmental constraints that restricts crop production worldwide. Introduction of stress-tolerant crop cultivars is the most auspicious way of surviving this constraint, and to produce these types of tolerant crops. Several bioengineering mechanisms involved in stress signaling are being adopted in this regard. One example of this kind of manipulation is the osmotic adjustment. The quarternary ammonium compound glycinebetaine (GB), also originally referred to as betaine is a methylated glycine derivative. Among the betaines, GB is the most abundant one in plants, which is mostly produced in response to dehydration caused by different abiotic stresses like drought, salinity, and extreme temperature. Glycinebetaine helps in decreased accumulation and detoxification of ROS, thereby restoring photosynthesis and reducing oxidative stress. It takes part in stabilizing membranes and macromolecules. It is also involved in the stabilization and protection of photosynthetic components, such as ribulose-1, 5-bisphosphate carboxylase/oxygenase, photosystem II and quarternary enzyme and protein complex structures under environmental stresses. Glycinebetaine was found to perform in chaperone-induced protein disaggregation. In addition, GB can confer stress tolerance in very low concentrations, and it acts in activating defense responsive genes with stress protection. Recently, field application of GB has also shown protective effects against environmental adversities increasing crop yield and quality. In this review, we will focus on the role of GB in conferring abiotic stress tolerance and the possible ways to engineer GB biosynthesis in plants.     

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.     

Polyamines and abiotic stress tolerance in plants

Environmental stresses including climate change, especially global warming, are severely affecting plant growth and productivity worldwide. It has been estimated that two-thirds of the yield potential of major crops are routinely lost due to the unfavorable environmental factors. On the other hand, the world population is estimated to reach about 10 billion by 2050, which will witness serious food shortages. Therefore, crops with enhanced vigour and high tolerance to various environmental factors should be developed to feed the increasing world population. Maintaining crop yields under adverse environmental stresses is probably the major challenge facing modern agriculture where polyamines can play important role. Polyamines (PAs)(putrescine, spermidine and spermine) are group of phytohormone-like aliphatic amine natural compounds with aliphatic nitrogen structure and present in almost all living organisms including plants. Evidences showed that polyamines are involved in many physiological processes, such as cell growth and development and respond to stress tolerance to various environmental factors. In many cases the relationship of plant stress tolerance was noted with the production of conjugated and bound polyamines as well as stimulation of polyamine oxidation. Therefore, genetic manipulation of crop plants with genes encoding enzymes of polyamine biosynthetic pathways may provide better stress tolerance to crop plants. Furthermore, the exogenous application of PAs is also another option for increasing the stress tolerance potential in plants. Here, we have described the synthesis and role of various polyamines in abiotic stress tolerance in plants.Key words: abiotic stress tolerance, putrescine, spermidine, spermine, polyamines     

植物水通道蛋白的干旱应答机制研究进展

干旱胁迫是严重影响全球作物生产的非生物胁迫之一,研究植物耐旱机制已成为一个重要领域。水通道蛋白是一类特异、高效转运水及其它小分子底物的膜通道蛋白,在植物中具有丰富的亚型,参与调节植物的水分吸收和运输。近10年来,水通道蛋白在植物不同生理过程中的作用,一直受到研究人员的关注,特别是在非生物胁迫方面,而研究表明水通道蛋白在干旱胁迫下对植物的耐旱性起着至关重要的作用,能维持细胞水分稳态和调控环境胁迫快速响应。水通道蛋白在植物耐旱过程中的调控机制及功能较复杂,而关于其应答机制和不同亚型功能性研究的报道甚少。该文综述了植物水通道蛋白的分类、结构、表达调控和活性调节,分别从植物水通道蛋白响应干旱表达调控机制、水通道蛋白基因表达的时空特异性、水通道蛋白基因的表达与蛋白丰度,水通道蛋白基因的耐旱转化四个方面阐明干旱胁迫下植物水通道蛋白的表达,重点阐述其参与植物干旱胁迫应答的作用机制,并提出水通道蛋白研究的主要方向。     

miRNAs: Major modulators for crop growth and development under abiotic stresses

Cumulatively, biotic and abiotic stresses of various magnitudes can decrease the production of crops by 70%. miRNAs have emerged as a genetic tool with enormous potential that can be exploited to understand stress tolerance at the molecular level and eventually regulate stress in crops. Plant miRNA targets frequently fit into diverse families of TFs that control the expression of genes related to a certain trait. As key machinery in gene regulatory networks, it is agreed that a broad understanding of miRNAs will greatly increase our understanding of plant responses to environmental stresses. miRNA-led stress regulatory networks are being considered as novel tools for the development of abiotic stress tolerance in crops. At this time, we need to expand our knowledge about the modulatory role of miRNAs during environmental fluctuations. It has become exceedingly clear that with increased understanding of the role of miRNAs during stress, the techniques for using miRNA-mediated gene regulation to enhance plant stress tolerance will become more effective and reliable. In this review we present: (1) miRNAs as a potential avenue for the modulation of abiotic stresses, and (2) summarize the research progress regarding plant responses to stress. Current progress is explained through discussion of the identification and validation of several miRNAs that enhance crop tolerance of salinity, drought, etc., while missing links on different aspects of miRNAs related to abiotic stress tolerance are noted.     

Interaction between PGPR and PGR for water conservation and plant growth attributes under drought condition

Drought is one of the key restraints to agricultural productivity worldwide and is expected to increase further. Drought stress accompanied by reduction in precipitation pose major challenges to future food safety. Strategies should be develop to enhance drought tolerance in crops like chickpea and wheat, in order to enhance their growth and yield. Drought tolerance strategies are costly and time consuming however, recent studies specify that plant growth promoting rhizobacteria (PGPR) and plant growth regulators (PGRs) can help plants to withstand under harsh environmental condition and enable plants to cope with drought stress. PGPR can act as biofertilizer and bioenhancer for different legumes and non-legumes. The use of PGPR and symbiotic microorganisms, may be valuable in developing strategies to assist water conservation in plants. The use of PGPR has been confirmed to be an ecologically sound way of enhancing crop yields by facilitating plant growth through direct or indirect mechanism. The mechanisms of PGPR for water conservation include secretion of exopolysaccharides, biofilm formation, alternation in phytohormone content, improvement in sugar concentration, enhancing availability of micro- and macronutrients and changes in plant functional traits. Similarly, plant growth regulators (PGRs) are specially noticed in actively growing tissues under stress conditions and have been associated in the control of cell division, embryogenesis, root formation, fruit development and ripening, and reactions to biotic and abiotic stresses and upholding water conservation status in plants. Previous studies also suggest that plant metabolites interact with plant physiology under stress condition and impart drought tolerance. Metabolites like, sugars, amino acids, organic acid and polyols play a key role in drought tolerance of crop plants grown under stress condition. It is concluded from the present study that PGRs in combination with PGPR consortium can be an effective formulation to promote plant growth and maintenance of plant turgidity under drought stress. This review is a compilation of the effect of drought stress on crop plants and described interactions between PGPR/PGRs and plant development, knowledge of water conservation and stress release strategies of PGPR and PGRs and the role of plant metabolites in drought tolerance of crop plants. This review also bridges the gaps that summarizes the mechanism of action of PGPR for drought tolerance of crop plants and sustainability of agriculture and applicability of these beneficial rhizobacteria in different agro-ecosystems under drought stress.     

Drought stress responses in crops

Among the effects of impending climate change, drought will have a profound impact on crop productivity in the future. Response to drought stress has been studied widely, and the model plant Arabidopsis has guided the studies on crop plants with genome sequence information viz., rice, wheat, maize and sorghum. Since the value of functions of genes, dynamics of pathways and interaction of networks for drought tolerance in plants can only be judged by evidence from field performance, this mini-review provides a research update focussing on the current developments on the response to drought in crop plants. Studies in Arabidopsis provide the basis for interpreting the available information in a systems biology perspective. In particular, the elucidation of the mechanism of drought stress response in crops is considered from evidence-based outputs emerging from recent omic studies in crops.     

Signal transduction and biotechnology in response to environmental stresses

Providing sufficient food to burgeoning population from the steadily shrinking arable land seems to be very difficult in near future and is one of the foremost challenges for plant scientists. In addition, there are several biotic and abiotic stresses which frequently encounter crop plants during various stages of life cycle, resulting in considerable yield losses. Environmental stresses, including drought, flooding, salinity, temperature (both low and high), high radiation, and xenobiotics induce toxicity, membrane damage, excessive reactive oxygen species (ROS) production, reduced photosynthesis, and altered nutrient acquisition. Several indigenous defence mechanisms (physiological and molecular) are triggered in plants on exposure to environmental cues. Enhancement of resistance of crop plants to environmental stresses has been the topic of prime interest for agriculturalists and plant scientists since long. Development of water and salinity stress-tolerant crops through genetic engineering provides an avenue towards the reclamation of farmlands that have been lost due to salinity and lack of irrigation water/rainfall. Understanding the complexity of stress tolerance mechanisms in orthodox or model plants at the genetic and molecular levels improves feasibility of enhancing tolerance of sensitive crop plants.     

Regulation in Plant Stress Tolerance by a Potential Plant Growth Regulator, 5-Aminolevulinic Acid

Exogenous application of different plant growth regulators is a well-recognized strategy to alleviate stress-induced adverse effects on different crop plants by regulating a variety of physiobiochemical processes such as photosynthesis, chlorophyll biosynthesis, nutrient uptake, antioxidant metabolism, and protein synthesis, which are directly or indirectly involved in the mechanism of stress tolerance. Of various environmental factors, salinity, drought, and extreme temperature (low or high) considerably diminish plant growth and yield by modulating endogenous levels as well as signaling pathways of plant hormones. Of various plant hormones/regulators, a potential plant growth regulator, 5-aminolevulinic acid (ALA), is known to be effective in counteracting the injurious effects of various abiotic stresses in plants. Until now the mechanisms behind ALA regulation of growth under stress have not been fully elucidated. It is also not yet clear how far growth and yield in different crops can be promoted by exogenous application of ALA and whether this ALA-induced growth and yield promotion is cost-effective. Thus, in this review we discuss at length the effects of ALA in regulating growth and development in plants under a variety of abiotic stress conditions, including salinity, drought, and temperature stress. Furthermore, advances in the functional and regulatory interactions of this plant growth regulator with plant stress tolerance, as well as the effective mode of exogenous application of ALA in inducing stress tolerance in plants are also comprehensively discussed in this review. In the future, overaccumulation of ALA in plants through manipulation of gene(s) could enhance plant stress tolerance. Thus, genetic manipulation of plants with the goal of attaining increased synthesis/accumulation of ALA and hence improved stress tolerance under stress conditions is an important area for research.     

植物抗旱和耐重金属基因工程研究进展

干旱和重金属污染严重影响植物的生长发育.植物耐逆相关基因的克隆和功能鉴定研究,为通过基因工程途径提高植物的抗逆性奠定了理论基础.水分亏缺、高盐、低温和重金属胁迫都能诱导LEA(late embryogenesis abundant protein)基因的表达.转基因研究表明,LEA蛋白具有抗旱保护作用、离子结合特性以及抗氧化活性;水孔蛋白存在于细胞膜和液泡膜上,在细胞乃至整个植物体水分吸收和运输过程中发挥重要作用.干旱和盐胁迫促进水孔蛋白基因转录物的积累.过量表达水孔蛋白可增强水分吸收和运输,提高植物的抗旱能力.金属转运蛋白参与重金属离子的吸收、运输和累积等过程.这些蛋白基因在改良草坪草植物的抗旱节水和耐重金属能力等方面具有潜在的应用价值.     

Understanding water deficit stress-induced changes in the basic metabolism of higher plants – biotechnologically and sustainably improving agriculture and the ecoenvironment in arid regions of the globe

Water is vital for plant growth, development and productivity. Permanent or temporary water deficit stress limits the growth and distribution of natural and artificial vegetation and the performance of cultivated plants (crops) more than any other environmental factor. Productive and sustainable agriculture necessitates growing plants (crops) in arid and semiarid regions with less input of precious resources such as fresh water. For a better understanding and rapid improvement of soil–water stress tolerance in these regions, especially in the water-wind eroded crossing region, it is very important to link physiological and biochemical studies to molecular work in genetically tractable model plants and important native plants, and further extending them to practical ecological restoration and efficient crop production. Although basic studies and practices aimed at improving soil water stress resistance and plant water use efficiency have been carried out for many years, the mechanisms involved at different scales are still not clear. Further understanding and manipulating soil–plant water relationships and soil–water stress tolerance at the scales of ecology, physiology and molecular biology can significantly improve plant productivity and environmental quality. Currently, post-genomics and metabolomics are very important in exploring anti-drought gene resources in various life forms, but modern agriculturally sustainable development must be combined with plant physiological measures in the field, on the basis of which post-genomics and metabolomics have further practical prospects. In this review, we discuss physiological and molecular insights and effects in basic plant metabolism, drought tolerance strategies under drought conditions in higher plants for sustainable agriculture and ecoenvironments in arid and semiarid areas of the world. We conclude that biological measures are the bases for the solutions to the issues relating to the different types of sustainable development.     

Structural evolution and function of stress associated proteins in regulating biotic and abiotic stress responses in plants

Adverse environmental conditions greatly influence crop production every year and threaten food security. Plants have a range of signaling networks to combat these stresses, in which several stress-responsive genes and regulatory proteins function together. One such important family of proteins, the Stress Associated Protein (SAP) family, has been identified as a novel regulator of multiple stresses. The SAPs possess a characteristic N-terminal A20 zinc-finger domain combined with either AN1 or C₂H₂ at the C-terminus. SAPs provide tolerance against various abiotic stresses, including cold, salt, drought, heavy metal, and wounding. The majority of SAPs are stress-inducible and have a function in conferring stress tolerance in transgenics. The role of SAPs in regulating biotic stress responses is a newly emerging field among researchers. SAPs interact with many other proteins to execute their functions; however, the detailed mechanism of these interactions needs to be elucidated. In this context, the present review provides a detailed view of the evolution and functions of SAPs in plants. The involvement in crosstalk between abiotic and biotic stress signaling pathways makes SAPs ideal targets to develop crops with tolerance against multiple stresses without any yield penalty. Altogether, we provide current knowledge on SAPs for investigating their role in stress response, which can further be exploited to develop climate-resilient crops through transgene-based, breeding-mediated, or genome-editing approaches.

     

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.