Plants are exposed to a variety of abiotic stresses in nature and exhibit unique and complex responses to these stresses depending on their degree of plasticity involving many morphological, cellular, anatomical, and physiological changes. Phytohormones are known to play vital roles in the ability of plants to acclimatize to varying environments, by mediating growth, development, source/sink transitions and nutrient allocation. These signal molecules are produced within the plant, and also referred as plant growth regulators. Although plant response to salinity depends on several factors; nevertheless, phytohormones are thought to be the most important endogenous substances that are critical in modulating physiological responses that eventually lead to adaptation to salinity. Response usually involves fluctuations in the levels of several phytohormones, which relates with changes in expression of genes involved in their biosynthesis and the responses they regulate. Present review described the potential role of different phytohormones and their balances against salinity stress and summarized the research progress regarding plant responses towards salinity at physiological and molecular levels. We emphasized the role of abscisic acid, indole acetic acid, cytokinins, gibberellic acid, salicylic acid, brassinosteroids, jasmonates, ethylene and triazoles in mediating plant responses and discussed their crosstalk at various baseline pathways transduced by these phytohormones under salinity. Current progress is exemplified by the identification and validation of several significant genes that enhanced crops tolerance to salinity, while missing links on different aspects of phytohormone related salinity tolerance are pointed out. Deciphering mechanisms by which plant perceives salinity and trigger the signal transduction cascades via phytohormones is vital to devise salinity related breeding and transgenic approaches.

Phytohormones and plant responses to salinity stress: a review

Plants are sessile beings, so the need of mechanisms to flee from unfavorable circumstances has provided the development of unique and sophisticated responses to environmental stresses. Depending on the degree of plasticity, many morphological, cellular, anatomical, and physiological changes occur in plants in response to abiotic stress. Phytohormones are small molecules that play critical roles in regulating plant growth and development, as well as stress tolerance to promote survival and acclimatize to varying environments. To congregate the challenges of salinity, temperature extremes, and osmotic stress, plants use their genetic mechanism and different adaptive and biological approaches for survival and high production. In the present attempt, we review the potential role of different phytohormones and plant growth-promoting rhizobacteria in abiotic stresses and summarize the research progress in plant responses to abiotic stresses at physiological and molecular levels. We emphasized the regulatory circuits of abscisic acid, indole acetic acid, cytokinins, gibberellic acid, salicylic acid, brassinosteroids, jasmonates, ethylene, and triazole on exposure to abiotic stresses. Current progress is exemplified by the identification and validation of several significant genes that enhanced crop tolerance to stress in the field. These findings will make the modification of hormone biosynthetic pathways for the transgenic plant generation with augmented abiotic stress tolerance and boosting crop productivity in the coming decades possible.

Potential role of phytohormones and plant growth-promoting rhizobacteria in abiotic stresses: consequences for changing environment.

Soil salinity and drought are major abiotic factors that limit crop growth and productivity worldwide. Indeed, soil salinity and drought disrupt the cellular ionic and osmotic balance. Although silicon (Si) is generally considered nonessential for plant growth and developments, Si uptake by plants can alleviate both biotic and abiotic stresses. Silicon application could therefore improve crop production under adverse climate and soil conditions. Several reports have reviewed the benefits of silicon application on crop growth, but the mechanisms of silicon action have not been systematically discussed. Here, we review recent advances on silicon uptake, transport, and accumulation in plants and how silicon alleviates salinity toxicity and drought stress. The major points are the following: (1) both passive and active silicon uptake may coexist in plants; (2) although silicon transporters have been identified in some plants, more silicon transporters remain to be identified, and the process of silicon transport needs further clarification; (3) the mechanisms for silicon-mediated tolerance of salinity and drought have been extensively investigated at both physiological and biochemical levels. The physiological aspects include increasing water uptake by roots, maintaining nutrient balance, decreasing water loss from leaves, and promoting photosynthetic rate. At the biochemical level, silicon may improve antioxidant defense abilities by increasing the activities of antioxidant enzymes and the contents of non enzymatic antioxidants; silicon may also contribute to osmotic adjustment and increase photosynthetic enzymatic activities; and (4) silicon can regulate the levels of endogenous plant hormones under stress conditions, whereas silicon involvement in signaling and regulation of gene expression related to increasing stress tolerance remains to be explored.

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Beneficial effects of silicon on salt and drought tolerance in plants

The combined effects of gibberellic acid and salinity on some antioxidant enzyme activities, plant growth parameters and nutritional status in maize plants

Salinity has plagued soil fertility and drastically affected growth and survival of glycophytes in irrigated regions of the world since the beginning of recorded history It is particularly common in arid and semi-arid areas where evapotranspiration exceeds annual precipitation, and where irrigation is therefore necessary to meet crop water needs Salt buildup in the soils and groundwater has threatened its productivity and sustainability Plant responses to salt stress include an array of changes at the molecular, biochemical and physiological levels Salt stress involves a water deficit induced by the salt concentration in the rhizosphere, resulting in disruption of homeostasis and ion distribution in the cell and denaturation of structural and functional proteins As a consequence, salinity stress often activates cell signaling pathways including those that lead to synthesis of osmotically active metabolites, specific proteins, and certain free radical scavenging enzymes that control ion and water flux and support scavenging of oxygen radicals or chaperones ROS detoxification forms an important defense against salt stress Legumes are a key component of sustainable agriculture and can offer many economic and environmental benefits if grown more widely in crop rotations because of their ability to fix nitrogen in the root nodules in a symbiotic interaction with soil rhizobia Due to their capacity to grow on nitrogen-poor soils, they can be efficiently used for improving saline soil fertility and help to reintroduce agriculture to these lands However, in legumes, salt stress imposes a significant limitation of productivity related to the adverse effects on the growth of the host plant, the root-nodule bacteria, symbiotic development and finally the nitrogen fixation capacity This paper reviews responses of legumes to salinity stress with emphasis on physiological and biochemical mechanisms of salt tolerance

Salinity and its effects on the functional biology of legumes

Application of NaCl (electrical conductivity 4.0 mS cm−1) resulted in about 52, 50 and 55 % reduction in total nitrogen contents in mung bean [Vigna radiata (L.) Wilczek] leaf, root and nodule, respectively. In nodule, nitrogenase activity was reduced by about 84 % under stress as compared with the control set. Glutamine synthetase activity was reduced by about 31, 16 and 23 %, glutamate oxoglutarate aminotransferase activity was reduced by 78, 57 and 42 % and glutamate dehydrogenase activity was reduced by 9, 8 and 42 % in leaf, root and nodule, respectively, under salt stress. The pretreatment with indole-3-acetic acid, gibberellic acid and kinetin, each ranging from 0.1 to 10 µM, in restoring the metabolic alterations imposed by NaCl salinity was investigated in mung bean. The three phytohormones used were able to overcome to variable extents the adverse effects of stress imposed by NaCl solution.

Effect of phytohormone pretreatment on nitrogen metabolism in Vigna radiata under salt stress

Efficiency of pretreatment as foliar spray of indole-3-acetic acid, gibberellic acid and kinetin, each ranging from 0.1 to 10.0 μM concentration, in restoring the metabolic alterations imposed by NaCl salinity was investigated in Vigna radiata (L.) Wilczek. Glycolate oxidase, superoxide dismutase, catalase and peroxidase activities increased under stress in leaves and roots also. Malondialdehyde content and total peroxide content also increased under stress. All the three hormones used were able to overcome to variable extents the adverse effects of stress imposed by NaCl to these parameters.

N. Chakrabarti

Papers

Effect of phytohormone pretreatment on nitrogen metabolism in Vigna radiata under salt stress

Alleviation of NaCl stress by pretreatment with phytohormones in Vigna radiata