Stressful environments such as salinity, drought, and high temperature (heat) cause alterations in a wide range of physiological, biochemical, and molecular processes in plants. Photosynthesis, the most fundamental and intricate physiological process in all green plants, is also severely affected in all its phases by such stresses. Since the mechanism of photosynthesis involves various components, including photosynthetic pigments and photosystems, the electron transport system, and CO2 reduction pathways, any damage at any level caused by a stress may reduce the overall photosynthetic capacity of a green plant. Details of the stress-induced damage and adverse effects on different types of pigments, photosystems, components of electron transport system, alterations in the activities of enzymes involved in the mechanism of photosynthesis, and changes in various gas exchange characteristics, particularly of agricultural plants, are considered in this review. In addition, we discussed also progress made during the last two decades in producing transgenic lines of different C3 crops with enhanced photosynthetic performance, which was reached by either the overexpression of C3 enzymes or transcription factors or the incorporation of genes encoding C4 enzymes into C3 plants. We also discussed critically a current, worldwide effort to identify signaling components, such as transcription factors and protein kinases, particularly mitogen-activated protein kinases (MAPKs) involved in stress adaptation in agricultural plants.

Photosynthesis under stressful environments: An overview

Plants are constantly challenged by various abiotic stresses that negatively affect growth and productivity worldwide. During the course of their evolution, plants have developed sophisticated mechanisms to recognize external signals allowing them to respond appropriately to environmental conditions, although the degree of adjustability or tolerance to specific stresses differs from species to species. Overproduction of reactive oxygen species (ROS) (hydrogen peroxide, H2O2; superoxide, O2ˉ˙; hydroxyl radical, OH. and singlet oxygen, 1O2) is enhanced under abiotic and/or biotic stresses, which can cause oxidative damage to plant macromolecules and cell structures, leading to inhibition of plant growth and development, or to death. Among the various ROS, freely diffusible and relatively long-lived H2O2 acts as a central player in stress signal transduction pathways. These pathways can then activate multiple acclamatory responses that reinforce resistance to various abiotic and biotic stressors. To utilize H2O2 as a signaling molecule, non-toxic levels must be maintained in a delicate balancing act between H2O2 production and scavenging. Several recent studies have demonstrated that the H2O2-priming can enhance abiotic stress tolerance by modulating ROS detoxification and by regulating multiple stress-responsive pathways and gene expression. Despite the importance of the H2O2-priming, little is known about how this process improves the tolerance of plants to stress. Understanding the mechanisms of H2O2-priming-induced abiotic stress tolerance will be valuable for identifying biotechnological strategies to improve abiotic stress tolerance in crop plants. This review is an overview of our current knowledge of the possible mechanisms associated with H2O2-induced abiotic oxidative stress tolerance in plants, with special reference to antioxidant metabolism.

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Hydrogen peroxide priming modulates abiotic oxidative stress tolerance: insights from ROS detoxification and scavenging.

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Physiological Plant Ecology Ecophysiology And Stress Physiology Of Functional Groups

Plants are frequently exposed to a plethora of unfavorable or even adverse environmental conditions, termed as abiotic stresses (such as salinity, drought, heat, cold, flooding, heavy metals, ozone, UV radiation, etc.) and thus they pose serious threats to the sustainability of crop yield. Soil salinity, one of the most severe abiotic stresses, limits the production of about 6 % of the world’s total land and 20 % of irrigated land (17 % of total cultivated areas) and negatively affects crop production worldwide. On the other hand, increased salinity of agricultural land is expected to have destructive global effects, resulting in up to 50 % land loss by the next couple of decades. The adverse effects of salinity have been ascribed mainly to an increase in sodium (Na+) and chloride (Cl–) ions and hence these ions produce the critical conditions for plant survival by intercepting different plant mechanisms. Both Na+ and Cl– produce many physiological disorders in plants but Cl– is the most dangerous. A plant’s response to salt stress depends on the genotype, developmental stage, as well as the intensity and duration of the stress. Increased salinity has diverse effects on the physiology of plants grown in saline conditions and in response to major factors like osmotic stress, ion-specificity, nutritional and hormonal imbalance, and oxidative damage. In addition to upper plant parts, salinity also affects root growth and physiology and their function in nutrient uptake. The outcome of these effects may cause the disorganization of cellular membranes, inhibit photosynthesis, generate toxic metabolites and decline nutrient absorption, ultimately leading to plant death. In recent decades, exogenous protectants such as osmoprotectants, phytohormones, signaling molecules, polyamines, antioxidants and various trace elements have been found effective in plants in mitigating the salt induced damages. These protectants showed the capacity to enhance the plants’ growth, yield as well as stress tolerance under salinity. In this chapter we attempt to summarize differential responses of plants to salinity with special reference to growth, physiology and yield. Further, we have discussed the progress made in using exogenous protectants to mitigate salt-induced damages in plants.

https://www.springer.com/cda/content/document/cda_downloaddocument/9781461447467-c1.pdf

Plant Response to Salt Stress and Role of Exogenous Protectants to Mitigate Salt-Induced Damages

The action of phytohormone producing bacteria and plant growth regulators on germination and seedling growth of wheat under saline conditions were studied. Seed dormancy enforced by salinity (100 mM NaCl) was substantially alleviated and the germination was promoted by gibberellin, auxin, zeatin, and ethephon from 54 to 97%. The IAA producing bacterial strains Pseudomonas aureantiaca TSAU22, Pseudomonas extremorientalis TSAU6 and Pseudomonas extremorientalis TSAU20 significantly increased seedling root growth up to 25% in non-salinated conditions and up to 52% at 100 mM NaCl, compared to control plants. It is concluded that growth regulators considerably alleviated salinity-induced dormancy of wheat seeds. The facts mentioned above make it possible to recommend root colonizing bacteria that produce phytohormone to alleviate salt stress of wheat grown under conditions of soil salinity.

Alleviation of salt stress by plant growth regulators and IAA producing bacteria in wheat.

https://cyberleninka.org/article/n/1152374.pdf

Physiological responses of three maize cultivars to drought stress and recovery.

Summary. In this study, effect of salinity with different osmotic potential on shoot length, total fresh and dry weight, amounts of organic (proline) and inorganic (K + and Na + ) substance of leaf tissue, the Na + /K + ratio, and leaf area, relative water content (RWC) and leaf osmolality in two maize (Zea mays L., var. intendata, C.6127 and DK.623) cultivars which are grown as a second yield in the Southeastern Anatolia Region (SAR) of Turkey, were investigated. Plants were grown for 30 days in the controlled growth room. Salinized culture solutions at different osmotic potential (0, -0.1, -0.3 and -0.5 MPa) prepared by adding varying amounts of NaCl and CaCl 2 to the main culture solution were applied to plants from the beginning of the germination. As a result the shoot length, total fresh and dry weight and the leaf area decreased, amounts of proline Na + , Na + /K + ratio and the leaf osmolality increased, but amounts of K + did not change significantly with increasing stress, and salt stress caused a similar decrease in leaf relative water content in both maize cultivars.

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The effect of salinity on some physiological parameters in two maize cultivars

The effect of NaCl (-0.1, -0.4 and -0.7 MPa) on some physiological parameters in six 23-day-old soya bean cultivars (Glycine max L. Merr. namely A 3935, CX-415, Mitchell, Nazhcan, SA 88 and Turksoy) at 25, 30 and 35 °C was investigated. Salt stress treatments caused a decline in the K + /Na + ratio, plant height, fresh and dry biomass of the shoot and an increase in the relative leakage ratio and the contents of proline and Na + at all temperatures. Effects of salt stress and temperature on Chl content, Chl a/b ratio (antenna size) and qN (heat dissipation in the antenna) varied greatly between cultivars and treatments; however, in all cases approximately the same qP value was observed. It indicates that the plants were able to maintain the balance between excitation pressure and electron transport activity. Pigment content and the quantum efficiency of photosystem II exhibited significant differences that depended on the cultivar, the salt concentration and temperature. The cultivars were relatively insensitive to salt stress at 30 °C however they were very sensitive both at 25 and 35 °C. Of the cultivars tested CX-415 and SA 88 were the best performers at 25 °C compared with SA 88 and Tiirksoy at 35 °C.

Effects of Salt Stress on Some Physiological and Photosynthetic Parameters at Three Different Temperatures in Six Soya Bean (Glycine max L. Merr.) Cultivars

The main objective of this study was to evaluate the effects of salt stress on the photosynthetic electron transport chain using two chickpea lines (Cicer arietinum L.) differing in their salt stress tolerance at the germination stage (AKN 87 and AKN 290). Two weeks after sowing, seedlings were exposed to salt stress for 2 weeks and irrigated with 200 ml of 200 mM NaCl every 2 days. The polyphasic OJIP fluorescence transient and the 820-nm transmission kinetics (photosystem I) were used to evaluate the effects of salt stress on the functionality of the photosynthetic electron transport chain. It was observed that a signature for salt stress was a combination of a higher J step (VJ), a smaller IP amplitude, and little or no effect on the primary quantum yield of PSII (φPo). We observed for AKN 290 a shorter leaf life cycle, which may represent a mechanism to cope with salt stress. For severely salt-stressed leaves, an inhibition of electron flow between the PQ pool and P700 was found. The data also suggest that the properties of electron flow beyond PSI are affected by salt stress.

Salt stress effects on the photosynthetic electron transport chain in two chickpea lines differing in their salt stress tolerance

Barley seedlings were pre-treated with 1 and 5 µM H2O2 for 2 d and then supplied with water or 150 mM NaCl for 4 and 7 d. Exogenous H2O2 alone had no effect on the proline, malondialdehyde (MDA) and H2O2 contents, decreased catalase (CAT) activity and had no effect on peroxidase (POX) activity. Three new superoxide dismutase (SOD) isoenzymes appeared in the leaves as a result of 1 µM H2O2 treatment. NaCl enhanced CAT and POX activity. SOD activity and isoenzyme patterns were changed due to H2O2 pre-treatment, NaCl stress and leaf ageing. In pre-treated seedlings the rate of 14CO2 fixation was higher and MDA, H2O2 and proline contents were lower in comparison to the seedlings subjected directly to NaCl stress. Cl− content in the leaves 4 and 7 d after NaCl supply increased considerably, but less in pre-treated plants. It was suggested that H2O2 metabolism is involved as a signal in the processes of barley salt tolerance.

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Papers

Physiological responses of three maize cultivars to drought stress and recovery.

The effect of salinity on some physiological parameters in two maize cultivars

Effects of Salt Stress on Some Physiological and Photosynthetic Parameters at Three Different Temperatures in Six Soya Bean (Glycine max L. Merr.) Cultivars

Salt stress effects on the photosynthetic electron transport chain in two chickpea lines differing in their salt stress tolerance

Pre-treatment with H2O2 induces salt tolerance in Barley seedlings