Loss of productivity in the major staple crops due to abiotic stress.
Abstract
Agricultural productivity world over is threatened by abiotic stress, intensifying food security issues. The plant hormones play a significant role in mitigating abiotic stresses, including drought stress, salinity stress, heat stress, and heavy metal stress, faced by the plants. Considerable research has been conducted to understand hormone-mediated abiotic stress responses in plants and the underlying biosynthetic and regulatory pathways. Deciphering these pathways would allow their manipulation in the laboratory and possible extension to the field. In the present chapter, an overview of the role plant hormones play in mitigating abiotic stress, the underlying mechanisms of their action, and the cross-talk between their signaling pathways to mitigate abiotic stress is presented.
Keywords
- abiotic stress
- plant hormones
- stress response
- stress mitigation
- plant productivity
1. Introduction
Plant hormones or phytohormones are biochemicals required for the normal growth and development of plants [1, 2, 3]. Plant hormones include auxins (IAA), gibberellins (GAs) cytokinins (CK), abscisic acid (ABA), ethylene (ET), besides jasmonates (JA), salicylic acid (SA), brassinosteroids (BR), strigolactones (SL), and nitric oxide (NO). Apart from their role in plant growth and development, hormones also mediate response to biotic (disease, pathogens, herbivores, etc.) and abiotic (drought, heat, salinity, heavy metals, etc.) stress [3, 4, 5, 6]. Hormones act at the site of their biosynthesis or some distance away from it [3, 6, 7, 8]. Hormone biosynthesis, distribution, and patterns of their signal transduction change under stress conditions [8, 9]. Ethylene and ABA play remarkable roles in regulating the abiotic stress response [8]. The exogenous supply of phytohormones also increases stress endurance in plants [10, 11]. Abiotic stress factors rarely occur individually, and many stresses produce the same effects at the cellular level with an overlap in the expression pattern of stress response genes [12]. In the current chapter, the hormone-mediated response of plants to the abiotic stress, including drought, heat, and salinity, is discussed.
1.1 What is abiotic stress?
Ecological factors favor plant growth at optimum levels and constitute stress at sub- or supra-optimal levels. Abiotic stress reduces crop productivity by about 50% (Table 1) [8, 29]. High temperatures lead to 20% decrease in the yield, low temperatures 7%, salinity 10%, drought 9% and other forms of stress cause 4% yield loss [30]. In grain crops, grain size, number, and dry weight are influenced by abiotic stress, especially if present during the reproductive phase [30]. Various aspects of plant growth as affected by abiotic stress are presented in Figure 1. Crop productivity may be reduced by 2.5–16% by a 1°C rise in seasonal temperature in tropical and subtropical regions [31]. The stress response depends on the genetic constitution and adaptive response of a plant [32].
Crop | Abiotic stress factor | Loss in productivity | References |
---|---|---|---|
Wheat | Drought | 27.5% | Zhang et al., [13] |
Temperature | 29–44% | Djanaguiraman et al., [14] | |
Salinity | 45% | Ali et al., [15] | |
Rice | Drought | 25.4% | Zhang et al., [13] |
Temperature | 3.2%* | Zhao et al., [16] | |
Salinity | 30–50% | Eynard et al., [17] | |
Maize | Drought | 5–15% | Campos et al., [18] |
Temperature | 7.4%* | Zhao et al., [16] | |
Salinity | 34% | Cucci et al., [19] | |
Chickpea | Drought | 45–69% | Nayyar et al., [20] |
Temperature | 39% | Devasirvatham et al., [21] | |
Salinity | 8–10% | Zawude and Shanko, [22] | |
Soybean | Drought | 46–71% | Samarah et al., [23] |
Temperature | 42–64% | Jumrani & Bhatia, [24] | |
Salinity | 66–86% | Bustingorri & Lavado [25] | |
Sunflower | Drought | 50% | Hussain et al., [26] |
Temperature | *6% | Rondanini et al., [27] | |
Salinity | 50% | El-Kader et al., [28] |
Table 1.
Estimates for 1°C rise in temperature.

Figure 1.
Schematic presentation of impact of abiotic stress on growth & development of plants.
1.2 Drought
Drought has been defined as “a period of abnormally dry weather sufficiently prolonged for the lack of water to cause a serious hydrologic imbalance in the affected area” [33]. Drought is one of the dominant factors diminishing crop productivity [34, 35]. Drought has been called as “one of the world’s extreme weather-related natural hazards” [35, 36]. It threatens the sustainability of agricultural systems around the world [37]. From 1994 to 2013, it represented 5% of all natural disasters and affected one billion people [35, 38]. All aspects of plant growth, including photosynthesis, protein synthesis, water relations, cell turgidity, membrane integrity, and nutrient uptake, are affected by drought [8, 39, 40]. It causes oxidative stress and damages the biological molecules, including DNA, proteins, and photosynthetic pigments. [8, 35, 41, 42, 43, 44, 45]. Plants synthesize a whole range of molecules as protection against drought stress, for example, proline, glycine betaine, soluble sugars (mannitol, sorbitol, and trehalose), polyamines, and proteins [37, 46].
ABA levels increase in plants under drought stress [47, 48], inducing the expression of ABA-dependent genes [6, 49]. ABA signaling leads to the closure of stomata, reducing transpiration [48]. Expression levels of
About 14 ABA receptor proteins mediate ABA signaling. Pyrabactin Resistance 1 (PYR1) and PYR1-like (PYL) regulatory elements undergo a conformational change after ABA binding and inactivate the clade A Serine/Threonine Protein Phosphatase 2C (PP2C) [48, 57, 58]. This in turn triggers the ABA signaling cascade by phosphorylation of serine/threonine kinases [48, 59]. Transcription of ABA-responsive genes is upregulated by binding of ABRE (ABA-Responsive Elements) to the ABRE-Binding Proteins (AREBs) or ABRE-Binding Factors (ABFs) [48, 60]. ABFs are activated by their ABA-mediated phosphorylation [48, 61].
In
ABA is negatively regulated by cytokinin receptor HKs, AHK2 (
1.3 Temperature
Temperature affects the distribution, phenology, and physiology of plants [89]. Temperature is increasing under dry as well as wet conditions in the changing global climate scenario [89, 90]. For 2081 – 2100, the IPCC has predicted average temperatures higher by 1.0°C to 1.8°C under very low, 2.1°C to 3.5°C under intermediate and 3.3°C to 5.7°C under very high GHG emission scenarios in comparison to 1850-1900 [91]. The crop productivity decreases by 6% for one degree rise in temperature beyond the optimum [8, 92]. Temperature stress causes accumulation of ROS, denaturation, misfolding, and aggregation of proteins, changes the membrane structure affecting permeability and raft distribution, besides its impact on leaf area, leaf retention, stomatal conductance, water potential, rate of transpiration, etc. [89, 90, 91, 93]. Photosynthetic capacity may be diminished or permanently damaged due to heat stress [91, 94].
Plants produce transcription factors, heat signaling proteins, and molecular chaperones to prevent protein misfolding and aggregation after heat shock (HS) [95, 96]. In response to HS, the endogenous ABA levels increase transiently increasing the antioxidant capacity [47, 97, 98], for example, by inducing RBOH-NADPH oxidases. Out of 10 different
Auxins play an important role in thermomorphogenesis [91, 98, 104]. Auxin biosynthesis genes
BRs increase the production of HSPs [91, 112] and regulate the heat-induced accumulation of proton–pumping ATPase and aquaporins [91, 113], besides inducing the expression and activity of ROS scavenging enzymes under heat stress [91, 114]. In tomatoes, BR treatment increases the expression of RBOH1 and apoplast H2O2 levels [115]. Interestingly, H2O2 activates MPK2, which in turn enhances RBOH1 expression [91, 116]. Heat stress causes the accumulation of BZR1 (Brassinazole-resistant 1), an important transcription factor in BR signaling, in the nucleus [110].
Ethylene is another hormone involved in heat stress tolerance. EIN2 and ER1 mutants have poor survival rates under heat stress [95, 97].
CKs play an important role in heat stress responses in plants [91, 119, 120]. They increase the activities of APX, SOD, and GP and also upregulate genes responsible for photosynthesis and carbohydrate metabolism under heat stress [91, 121]. CK oxidase/dehydrogenase inhibitors improve heat stress tolerance [91, 122]. Heat stress tolerance is also increased in plants with ectopic expression of isopentenyl transferase (ipt) from
In
GA biosynthesis and accumulation increase under elevated temperatures in Carrizo citrange seedlings, wheat, and soybean hypocotyl [98, 129]. PIF4 upregulates the
SA biosynthesis is suppressed at higher temperatures in tobacco after TMV infection and in
1.4 Salinity
Soils with electric conductivity higher than 4 dS/m at 25°C are classified as saline [134]. Salinity has affected more than 800 million hectares of land globally, decreasing potential agricultural land by 1–2% per year [8, 135]. More than 50% of the land in developing countries, particularly that falling in the arid region, is affected by salinity, causing yield losses to the tune of 40%, for example, in the case of wheat [8, 136]. It decreases the quantity as well as the quality of the produce [8, 137]. Salinity impairs water uptake and causes ion toxicity, osmotic stress, nutrient deficiency, and oxidative stress [138]. Salt stress causes physiological drought impairing protein and photosynthesis [8, 137]. Changes in the intracellular Ca2+ levels, excess Na+, and ROS accumulation are the signals that trigger the salt stress response [139].
Ethylene is the major hormone in the salt stress response [138]. The levels of ET as well as its precursor ACC (1-aminocyclopropane-1-carboxylate) increase under salt stress [138, 140]. While salt tolerance can be increased by the application of ET or its precursor ACC [141, 142], inhibition of ET synthesis or signaling may increase salt sensitivity [138]. Ethylene signaling involves five ethylene receptors [ETR1 (Ethylene Response 1), ERS1 (Ethylene Response Sensor 1), ETR2, EIN4 (Ethylene Insensitive 4), and ERS2], a protein kinase, CTR1 (Constitutive Triple Response 1—a negative regulator) and a key positive regulator EIN2, which signals primary transcription factors EIN3, EIL1 (Ethylene Insensitive Like 1) and EIL2 and many downstream ethylene response factors. Osmotic stress, induced by many abiotic stresses, including salinity, suppresses the expression of
200 mM NaCl induces the expression of
An array of stress-responsive genes is regulated by ABA [153]. ABA coordinates with ET in mediating salt stress. On exposure to salt, many genes involved in ABA biosynthesis, including ZEP, AAO, and MCSU, are stimulated through Ca2+ −dependent phosphorylation events and their downstream signaling pathways [153, 154]. Increased ABA levels have been reported in many plants, including
The information available on the mechanism of salt stress response via auxins is scarce [153].
2. Conclusions
Plant hormones play an important role in the growth and development of plants and also represent an important line of defense against abiotic stress. Hormones change the pattern of growth to enable the plants to withstand stress. The plant stress response involves many hormones, their downstream response factors, associated gene networks, and transcription factors. The crosstalk between hormones and their synergistic or antagonistic interactions play central role in phytohormone-mediated abiotic stress tolerance [165]. Understanding the molecular level interaction between elements of different pathways controlling stress response is critical to allow their manipulation to improve stress tolerance. This is important, as the diversity, duration, and intensity of abiotic stresses are increasing in the changing global climate scenario. Plant hormones are an important target for better management of abiotic stress, especially, in view of the limited success of conventional breeding techniques in dealing with it. Phytohormone pathways and the intermediaries therein can go a long way in the production of climate-resilient crops.
New technologies to bioengineer plants have proven useful in achieving this end; examples include soybean [166], maize [167], rice [168], and potato [169]. Techniques including transcriptome analysis, next-generation sequencing analysis, transgenic plants, genome editing, etc. are being used to identify the hormone-mediated regulatory mechanisms of the plant stress response. Transcriptome analysis using microarrays, a survey of transcriptome profiles, and levels of microRNAs in plants under stress using RNA-seq have helped understand the mechanism of stress tolerance in plants [170]. With genome editing technology, genomes can now be modified in a site-specific manner using specifically designed endonucleases like zinc finger nucleases (ZFN) or TAL effector nucleases (TALEN; [49, 171]) and the CRISPR/CAS system [49, 172].
In a nutshell, new pathways are already emerging. However, the complex interactions between the hormones and their ability to regulate a wide array of plant developmental and physiological processes complicate teasing out the effect of an individual hormone. Lack of information about the tissue-specific stress response and genetic plasticity as well as the extreme complexity of thresholds for different stress responses makes mechanistic understanding of abiotic stress tolerance difficult [173]. In order to better understand the hormone mediated abiotic stress response, the future research should focus on identifying the antagonistic and synergistic interactions between various hormones and the critical regulatory junctures in the hormone crosstalk.
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