Identification of Different Stress Responsive Genes induced in Crops for Climate Changes: A Review

Author:

Rajendra Prasad Meena1*, Deen Dayal Bairwa1, Vikas Sharma2, Sanjiv Kumar1, Satyesh Raj Anand3, Harkesh Kumar Balai1, Vijay Daneva1 and Ashish Kumar Sharma1

Journal Name: Biological Forum – An International Journal, 16(5): 06-17, 2024

Address:

1Department of Agriculture, Jagannath University, Chaksu, Jaipur (Rajasthan), India.

2Swami Keshwanand Rajasthan Agricultural University, Bikaner (Rajasthan), India.

3Faculty of Engineering and Technology, Jagannath University, Chaksu, Jaipur (Rajasthan), India.

(Corresponding author: Rajendra Prasad Meena*)

DOI: -

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Abstract

Numerous environmental stresses affect plants, lowering and restricting the productivity of crops used in agriculture. Plants are subject to two ecological stresses: biotic and abiotic stress. Major crop plants are lost due to abiotic stress, which includes radiation, salinity, floods, droughts, temperature extremes and heavy metals. Biotic stress can result in attacks by various pathogens, including bacteria, fungi, oomycetes, nematodes and herbivores. Due to their sessile nature, plants cannot avoid these environmental cues. Plants have evolved various defense mechanisms to deal with these dangers from biotic and abiotic stresses. Ecological stresses have a disastrous effect on the growth and yield of plants in the field. Recent research has shown that plants' responses to combinations of two or more stress conditions are unique and cannot be directly extrapolated from the responses of plants to each of the different stresses applied individually. The field environment differs significantly from the controlled conditions used in laboratory studies and frequently involves the simultaneous exposure of plants to multiple abiotic and biotic stresses conditions.

Keywords

Environmental stresses, temperature, fungi, drought, radiation, bacteria, nematodes and salinity.

Introduction

The increasing risks that global warming to crop growth and yield due to biotic and abiotic stresses make it a cause for concern worldwide (Ramegowda and Senthil Kumar 2015). Abiotic stresses that adversely affect productivity in agriculture include salinity, drought and heat and cold waves. Currently, abiotic stresses like heat, cold, drought and salinity account for more than 60% of yield losses in all major crops (Reis et al., 2012). The beneficial goal of "food for all," set by nations worldwide is made more difficult by the growing global population. By 2050, the estimated 60% increase in cereal production is undoubtedly insufficient to feed the world's population of approximately 9.7 billion people. Achieving sustainable growth in food production for the growing population necessitates making the best use of the resources at hand. Expanding the productivity of the world's harsher environments and raising agricultural productivity is critical given the exponential growth in global population. Regrettably, the most of land with potential for agriculture is located in arid regions. Several studies have revealed that India's agriculture, food security and water availability have all been negatively impacted by climate change (Burney et al., 2014). One of the main consequences of climate change is an increase in air temperature, which causes yields to drop sharply and may lead to significant changes in cropping patterns. It is well known that high temperatures cause plants to respire more quickly, which may lessen the benefits of higher photosynthetic rates brought on by higher CO2 concentrations (Bagley et al., 2015). According to Hof et al., (2015), an increase in the average temperature is foreseen to increase the frequency of illnesses and pest damage as well as evaporation. This could lead to a need for irrigation water, a resource that is already running out globally (Fader et al., 2016). Consequently, plants have been observed to experience an increase in drought or low moisture stress as a result of rising air temperatures, which poses a significant challenge to crop productivity globally. The average yields of major crops are declining due to the rapid increase in desertification worldwide (Bray, 1997). Reduced moisture stress lowers crop yields in arid and semi-arid areas. Plants under drought stress undergo various physiological, biochemical and molecular changes. Both soil and plant water potential declines under drought stress. Drought stress results in decreased water potential and reduced turgor pressure, stomata closure, leaf growth and rate of photosynthesis (Monte, 1986; Ozturk, 1998). Low water stress is also known as drought stress, it occurs when a plant experiences a more water loss through transpiration than it takes up through its roots, leaving the plant in a state of water deficit. Drought tolerance in agriculture refers to a crop plant's capacity to endure and maintain productivity under prolonged environmental conditions of water deficit with loss (Mitra, 2001). Various morphological, physiological and biochemical characteristics results in drought tolerance. According to Ingram et al. (1996); Zandalinas et al. (2017), Plants alterations may relate to gene expression, metabolic pathways, osmotic adjustments, repair systems, etc. The mechanisms underlying plants' ability to withstand drought have been the subject of numerous attempts in the fields of molecular biology and genomics. Numerous transcriptionally-responsive genes to drought stress have been found and confirmed (Growth et al., 2010; Joshi et al., 2016). Plants respond differently to drought stress, these changes are essential in determining a plant's survival, distribution throughout its range and yield. While most crops are sensitive to water shortages, especially in the flowering to seed development stage, some crops grown in semi-arid and arid regions, like pigeon pea, pearl millet and sorghum can tolerate drought. This suggests that these crops have developed numerous interconnected strategies and a tolerance mechanism to survive in drought-prone environments. It is challenging to characterize the phenotypic and physiological parameters required for choosing the improved crop under drought stress due to the wide variation in the duration and severity of drought stress and the lack of knowledge regarding the complexity of drought stress. Genomics techniques such as genome sequencing have partially addressed this challenge, which is crucial for identifying genes linked to drought tolerance in many crops and creating markers connected to drought stress. These days, many significant crop genomes have already been sequenced, providing a wealth of opportunities to comprehend the molecular mechanism underlying the crops' ability to withstand abiotic stress. On the other hand, not much genomic research has been done on orphan crops like pearl millet. The current emphasis is on characterizing these orphan plant species transcriptomes to understand the composition and function of various genes involved in the response to drought stress. The most practical way to investigate the differential expression of genes is to create cDNA-based EST (Expressed Sequence Tags) libraries. The EST sequence data constitutes a valuable resource that may be applied to various genomic research projects. Furthermore, according to Mishra et al. (2007), it serves as a platform for comprehending the mechanisms underlying plant adaptation to multiple stresses. Given the growing drought conditions, we must create food crop genotypes resistant to abiotic stress conditions. The naturally occurring variations of our major crops are becoming a limited gene pool, which is restricting breeding strategies for crop improvement. For this reason, we must search the varied natural habitat for genetic resources to find novel genes or allelic variations of already-known genes. Recombinant DNA technology can be useful for transfer genes from such tolerant crops to cereal crops, creating transgenics that are resistant to drought. As a member of the Poaceae family, pearl millet may be a more suitable substitute than genes from other dicots or extremophiles for creating transgenic cereal crops resistant to drought stress.

Biotic stresses. Stress commonly referred to as "biotic stress" is the result of an organism being harmed by other living things, including weeds, native or cultivated plants, bacteria, viruses, fungi, parasites and both beneficial and harmful insects. An organism's ability to withstand specific stresses and the climate in which it lives determine the kinds of biotic stresses that it faces. The term "biotic stress" is still widely used and researchers dealing with it encounter numerous obstacles. For example, regulating biotic stress in an experimental setting is more complex than controlling abiotic stress (Flynn, 2003). Agricultural research has made biotic stresses a primary focus because of the substantial losses caused by cash crops deteriorates. Plant yield and biotic stress influences both practical development and economic decisions. Plant-stressor convolution, population dynamics and ecosystem nutrient cycling are all impacted by the effects of biotic injury on crop yield (Peterson et al., 2001).

Inducible defense responses to insect herbivores

A plant must be distinguished between abiotic and biotic stress to protect itself against it plant reacts to a herbivore by recognizing specific chemicals prevalent in the herbivore's saliva. Herbivore-associated molecular patterns (HAMPs) or elicitors are the substances that cause a reaction in plants. By triggering signaling pathways throughout the plant, these HAMPs enable the plant to minimize damage to other regions and initiate its defense mechanism. By leading to signaling pathways throughout the plant, these HAMPs enable the plant to minimize damage to other regions and initiate its defense mechanism. Similar to aphids, phloem feeders do not inflict significant mechanical harm on plants; however, they are nonetheless considered pests and have the potential reduce crop yields significantly. To defend themselves against phloem feeders, plants have evolved a defense mechanism using the salicylic acid pathway, which is also used in infection stress. Plants target an insect's digestive system more directly. Plants use inhibitors of proteinase to achieve these. While these proteinase inhibitors enter an insect's digestive system, they attach firmly and precisely to the active site of enzymes that hydrolyze proteins, like trypsin and chymotrypsin, preventing the digestion of proteins (Taiz et al., 2015).

Inducible defense responses to pathogens. Despite lacking an immune system or circulatory system like those found in animals, plants can recognize non-self-signals and use them to detect invaders. Identifying microorganism-associated molecular patterns, or MAMPs, is frequently a plant's first line of defense against microorganisms at the cell surface (Spol et al., 2012). MAMPs consist of endotoxins on bacterial cell membranes and nucleic acids shared by viruses that specific pattern-recognition receptors recognize. Utilizing plant immune receptors to identify effector molecules that infections release into plant cells is another technique for detection. Effector-triggered immunity (ETI), a subset of the innate immune response, is activated upon detecting these signals in infected cells (Tsuda et al., 2010). It is known that pathogenic infection causes an increase in the synthesis of salicylic acid (SA). Plant resistance to biotrophic and hemibiotrophic pathogens is ultimately increased by producing of pathogenesis related (PR) genes, in response to elevated SA levels. According to Bari et al., (2009), the ubiquitination of jasmonate ZIM domains (JAZ) proteins, which block JA signaling, is a physiological reaction to increased JA production. This leads to the degradation of the JAZ proteins and an increase in JA-activated defense genes.

Abiotic Stresses. During their growth period, crop plants are subjected to various abiotic stresses, including but not limited to drought, salinity, extreme temperatures, submergence and water logging, all of which have a negative impact on their overall growth and development (Zhang et al., 2010). These stresses can cause the average yields of most crops to drop by more than half. According to estimates, the global population could reach nine billion people by 2050, necessitating a rise in crop yields (Pennisi, 2008; Nakashima et al., 2014). To adapt to these environmental stresses, plants change the morphological and biochemical phenotypes that affect their physiological responses and ultimately aiding their survival. Under various ecological circumstances, these stresses force plants to adapt in multiple ways, which serve as the foundation for speciation. According to the Intergovernmental Panel on Climate Change (IPCC), abiotic stresses will rise shortly due to global climate change. Abiotic stresses come in many forms and they can be applied to plants simultaneously or at different stages of their development (Tester and Bacic 2005; Hirayama and Shinozaki 2010). Plant cells absorb these stress signals at the membrane or cytoplasmic level. Plant cells absorb these stress signals through a wide range of sensors and receptors at the membrane or cytoplasmic level. They then convert these signals into a variety of signal transduction cascades that induce both primary and secondary stress responsive genes (heat shock proteins (HSPs), protective proteins, ROS scavenging enzymes, antioxidants, compatible osmolytes, etc.), which result in stress adaptation (Shinozaki and Yamaguchi-Shinozaki, 2000).In response to these stresses, plants exhibit dynamic responses involving intricate cross-talk at multiple regulatory levels (Krasensky and Jonak 2012). Changes in the transcriptome, proteome and metabolome levels are associated with plant responses. These reactions are unique to multiple stressors as opposed to single ones and every type of stress triggers a distinct gene repertoire associated with the particular environmental circumstances experienced. The agricultural sector has emphasized developing stress-resistant crops modified to grow in variable environmental conditions and provide higher plant productivity to make up for productivity losses in crops caused by stress conditions. Stress-responsive genes are the best options to giving crops stress tolerance. Identifying QTLs related to abiotic stress and their application in marker-assisted breeding and transgenic developments are underway in various crop varieties. Functional genomics has been instrumental in analyzing the genes linked to different environmental stressors, enabling the selection of more resilient and robust crops in the future.

Drought Stress. Plants under drought stress due to a lack of water. The average metabolic process of crop plants is hampered and yield is decreases when water is unavailable to them for extended periods of time. Plants attempt to survive in these circumstances by sustaining the barest minimum of physiological activity, which entails controlling thousands of genes and different metabolic pathways (Passioura, 1997; Mitra, 2001; Luo, 2010).

Drought stress responses in crop plants. Plants undergo various physio-biochemical changes in response to drought (Table 1). The physiological changes in plants include decreased leaf area, rolling, drying and increased root growth. The biochemical changes include producing reactive oxygen species (ROS) that disrupt cellular homeostasis, accumulation of free radicals and electrolytic leakage (EL). The buildup of reactive oxygen species (ROS) leads to the breakdown of different biomolecules found in cells, including proteins, lipids, nucleic acids and pigments. This membrane damage ultimately compromises the cell's viability (Bartels and Sunkar 2005). Additionally, hormones are crucial in a variety of abiotic stress situations. Among other hormones, ABA is recognized to be important in responding to a variety of abiotic stressors, including salinity, cold and drought (Zhu, 2002). According to Seki et al. (2002), ABA treatment can also induce many genes in response to abiotic stresses, offering direct evidence of its role in these environmental stresses. Drought stress is directly related to ABA; it causes stomata to close to prevent water loss through transpiration and lowers the rate of photosynthesis in order to raise plants' water-use efficiency (WUE). In addition to these reactions, there are molecular reactions, which comprise signal transduction, perception and modification of gene expression and metabolic alterations that are communicated. In addition to producing vital metabolic proteins that shield cells from stress, the changed gene expressions brought on by drought stress also control the downstream signal transduction pathways.

Drought stress negatively impacts vegetative growth (Tripathy et al., 2000). It has been reported that soybeans matured eight days earlier than usual due to a reduced vegetative growth. Extended drought stress further restricted its capacity to form seeds (Twidwell, 2002). Under stressful situations, there was a significant decrease in relative water content (RWC) and an increase in free proline content. One of the most often produced compatible osmolytes in response to water stress conditions is proline. Under stressful situations, the genotype showed elevated activity of peroxidase (POD) and superoxide dismutase (SOD) and decreased accumulation of catalase (CAT). According to the data, proline content and the antioxidant defense mechanism activated during water stress are related. PEG 6000-induced drought stress reduced pearl millet germination percentage and shoot growth while slightly lengthening the roots. Plants' adaptive response to drought stress was reflected in an increase in root length (Leila Rad, 2007). Several ROS-scavenging enzymes are also activated during drought stress. Ascorbate peroxidase (APX) activity was higher in mildly stressed plants than in control plants (Sharma and Ghildiyal 2005). However, at higher levels of drought stress, APX activity declined in the initial set of plants.

Table 1: Higher plants 'Physiological, biochemical and molecular responses under moisture deficit stress conditions (source: Reddy et al., 2004).

Physiological responses

Biochemical responses

Molecular responses

- Recognition of root

- Transcient decrease in

- Stress responsive gene

signal

Photochemical efficiency

expression

- Loss of turgor and

- Decreased efficiency of

- Increased expression in

osmotic   adjustment

Rubisco

ABA biosynthetic genes

- Reduced leaf Ψw

- Accumulation of stress

- Expression of ABA

- Decrease in stomatal

metabolites like MDHA,

responsive  genes,

conductance to CO2

Glutathione, Proline,

- Synthesis of specific

- Reduced internal CO2

Glycine betaine

proteins  like Late

concentration

Polyamines and α-

embryogenesis abundant,

- Decline in net

tocopherol

- Desiccation stress protein,

photosynthesis

- Increased in antioxidants

dehydrins, etc.

- Reduced growth rates.

- Reduced ROS accumulation

- Drought stresses tolerance



Mechanism of drought tolerance in plants. Plant drought resistance is stage-specific and linked to several physiological, molecular and phenological processes. As a result, it is difficult to thoroughly and accurately analyze a particular plant species' entire drought resistance mechanism. A complex trait, drought resistance is linked to multiple physiological characteristics, Plants develop variety of various defense mechanisms, to combat drought stress, such tolerance avoidance and escape (Levitt, 1972). Plants can finish their life cycle during a drought before experiencing extreme stress (Mooney et al., 1987). To avoid having the growing season coincide with a localized seasonal or environmental drought, it refers to the readjustment of the life cycle or growth and developmental timings (Mitra, 2001; Manavalan et al., 2009). Conversely, drought avoidance minimizes water loss and maximizes water uptake to avoid drought conditions (Chaves et al., 2003). The ability of a plant to generate a commercial product under drought stress with the least amount of loss is known as drought tolerance. It is a multifaceted characteristic that involves adaptations on many levels, ranging from physiological and biochemical processes to plant morphology and anatomical structures (Blum, 2002). Numerous genes undergo transcriptional reprogramming in conjunction with drought tolerance. According to Joshi et al., (2016), transcription factors are significant in controlling the expression of genes that respond to stress and act downstream during stressful situations. The three components of drought resistance in plants, drought avoidance and drought tolerance are the two main mechanisms for drought resistance (Yue et al., 2006). Deciphering the genetic basis of drought tolerance in plants remains challenging due to its quantitative nature and multiple governing genes (Price et al., 2002).

Approaches for the development of stress tolerant crops 

Conventional breeding approach. Drought-tolerant crops have been developed through conventional and molecular breeding techniques and some commercially viable hybrid cultivars and inbred lines with drought tolerance have been developed and released (Scott et al., 1986). However, the main limitations of this approach, are the complex nature of the trait, the absence of desired genetic variations in the available germplasm pool and the need for adequate screening facilities for genotyping a large breeding population.

Transgenic approaches. Using the concepts of genetic engineering, the transgenic approach transfers genes from genotypes resistant to drought to genotypes susceptible to it. However, it requires thorough knowledge of the genes' roles concerning to drought stress. One of the main drawbacks of this approach is the selection of the major candidate gene for the desired trait, since drought tolerance involves multiple genes. Drought-tolerant crops have recently been developed in various plant species through genes coding for stress-responsive transcription factors and stress-resistant proteins (Table 2). Furthermore, current research focuses on clarifying and characterizing these critical factors and their mechanisms of action because the factors that confer enhanced drought tolerance in higher plants still need to be better understood. A more profound comprehension of the genes involved in the drought tolerance mechanism is required to genetically modify plants. The process of mapping QTLs that confer drought tolerance will make it easier to use marker-assisted breeding to develop drought-tolerant varieties.




Table 2: Major abiotic stress tolerant genes characterized in various crop species.

Source of gene

Name of gene

Transformed receptor

Characteristics

Reference


A. thaliana


DREB2A

Saccharum sp.

Drought tolerant

Reis et al. (2014)

MYB96

Camelina sativa

Heat tolerance

Lee et al. (2014)

Rab7

A. thaliana

Drought and salt tolerant

Mazel et al. (2004)

SAP5

Gossypium hirsutum

Drought tolerant

Hozain et al. (2012)

HARDY

O. sativa

Drought tolerant

Karaba et al.(2007)

Hsp17.6A

A. thaliana

Drought and salt tolerant

Sun et al. (2001)

T. aestivum

NAC2a

Nicotiana

tabacum

Drought tolerant

Tang et al. (2012)

PIMP1

T. aestivum

Drought tolerant

Zhang et al. (2012)

WRKY1 and WRKY33

A. thaliana

Drought and heat tolerance

He et al. (2016)

ASR1

N. tabacum

Drought tolerance

Hu et al. (2013)

ASR5

A. thaliana

Drought tolerant

Li et al. (2017)

AREB1and ABF3

A. thaliana

Drought tolerant

Yoshida et al. (2010)

DREB1A

O. sativa

Drought tolerant

Datta et al. (2012)

ABF3

A. thaliana

Drought tolerant

Oh et al. (2005)

O. sativa

bZIP46

O. sativa

Drought and salt tolerant

Tang et al. (2012)

MYB3R-2

A. thaliana

Drought and salt tolerant

Dai et al. (2007)

BiP

A. thaliana

Drought tolerant

Valente et al. (2009)

G. max


P5CR

A. thaliana

Drought, salt and Heat tolerant

De Ronde et al. (2004)

NF-YB2

A. thaliana

Drought tolerant

Nelson et al. (2007)

Z. mays


NPK1

A. thaliana

Drought tolerant

Shou et al. (2004)

NHX3

T. aestivum

Drought tolerant

Zörb et al. (2004)

SacB

A. thaliana

Drought tolerant

Pilon-Smits et al. (1995)

N. tabacum


APX

A. thaliana

Drought and salt tolerant

Badawi et al. (2004)

HVA1

Morusindica

Drought and salt tolerant

Checker et al. (2012)

Hordeum vulgare

WRKY5

A. thaliana

Drought tolerant

Ma et al. (2014)

Leymus chinensis

LAS

A. thaliana

Drought and salt tolerant

Yang et al. (2011)

Brassica napus

ASR4

A. thaliana

Drought tolerant

Li et al. (2017)

Setaria italica

IF4A

Arachishy

pogaea 


Drought and salt tolerant

Rao et al. (2017)

Pennisetum glaucum

ASR1

E. coli

Drought tolerant

Padaria et al. (2016)



Methods to identify stress tolerant genes. To ascertain the differential expression analysis of genes under various conditions and periods, a number of molecular techniques are available. Among these methods are microarrays, serial analysis of gene expression (SAGE), subtractive hybridization, suppression subtractive hybridization (SSH), differential display reverse transcription PCR (DDRT-PCR), etc. In order to identify P. glaucum stress-related genes, James et al. (2015); Choudhary et al. (2015) employed the SSH technique under two distinct stress conditions heat and drought. ASR protein, APX, glyoxalase, Rab7 and aspartic proteinase were among the genes that Choudhary et al. (2015) found to be differentially expressed when they used PEG 30% for varying lengths of time to impose drought stress. These findings were then confirmed by qRT-PCR.

Genes induced during drought stress. It's crucial to comprehend the molecular subtleties of drought response in order to impart drought tolerance in crops. Using different approaches to gene mining, remarkable information about the types of drought responsive genes induced in other crop species has been obtained. Numerous details regarding the important genes driving the drought stress response have been made available by differential analysis at the transcriptome, proteome and metabolome levels. The synthesis of novel metabolic proteins, including osmoprotectants, heat shock proteins (HSPs), Late Embryogenesis Abundant (LEA) proteins, detoxifying enzymes, antioxidants, water channels like aquaporins, ion pumps and regulatory proteins, which primarily consist of TFs (AREB, AP2/ERF, NAC, bZIP, MYC, and MYB), is either facilitated by these important genes. Protein phosphatases and kinases participate in the synchronizing of signal transduction pathways (Wani et al., 2013). Important genes for the production of metabolic proteins, Sugars (fructose, fructan), sugar alcohols (mannitol, sorbitol) and zwitterionic substances (proline, glycine, betaine) are examples of compatible osmolytes. Due to their high water solubility, these build up in cells at higher concentrations under various water deficit conditions. Higher concentrations of these metabolites within the cell aid plants in coping with water deficit stress in two ways: they increase water retention or osmotic adjustment, which keeps cells hydrated and they stabilize cellular molecules by acting as osmoprotectants, which protect them from the damaging effects of ionic stress. According to Delauney and Verma (1993), proline is the most widely distributed compatible solute-small hydrophilic organic molecules that build up to high concentrations-in water-stressed plants and numerous other organisms. Proline's enhanced biosynthesis in dehydrated plants and the inhibition of its degradation both contribute to its higher accumulation. In rice under water stress, the enzyme Δ1-pyrroline-5-carboxylate synthase (P5CS) was induced (Zhu et al., 1998). While the suppression of P5CR resulted in increased sensitivity to various abiotic stresses, higher proline content encouraged better growth under conditions of water scarcity (De Ronde et al., 2000). According to additional data, Proline may help the DREB1 pathway facilitate drought adaptation. As an illustration, it has been demonstrated that transgenic rice plants that overexpress OsDREB1 or AtDREB1 (Ito et al., 2006) accumulate more proline than wild-type plants under both normal and water-deficit conditions, and they also exhibit superior adaptation to water stress conditions. Glycine Betaine (GB): Several plant families, including the Composite, Chenopodiaceae, and Gramineae, have reported on the function of glycine betaine in response to drought and salinity stress. In these and many other halotolerant plants, GB builds up in the plastids and chloroplasts (Chen and Murata, 2008). Higher levels of GB accumulation were observed in transgenic maize plants transformed with the choline dehydrogenase coding bet A gene, which regulates drought stress in the field, according to Quan et al. (2004). After three weeks of drought stress, transgenic plants produced 10-23% more grain per plant than untransformed plants, while control plants' reproductive development was significantly hindered. Under drought stress, BetA-transgenic cotton outperformed the wild-type controls in several physiological parameters (Lv et al., 2007). Trehalose is a disaccharide that does not reduce and is crucial in the abiotic stress. It prevents denaturation of the protein and other biomolecules. Two essential enzymes, trehalose-6-phosphate synthase (TPS) and trehalose-6-phosphate phosphatase (TPP), catalyze trehalose biosynthesisin plants. However, the majority of higher plant species only accumulate trace amounts of trehalose, except highly desiccation-tolerant resurrection plants. When various abiotic stresses, such as dehydration, cold and salinity occur LEA proteins are protective and accumulate at higher concentrations. These proteins are often produced in developing embryos during the desiccation of seeds and they are also induced in stressed vegetative tissues. The hydrophilic LEA proteins are primarily responsible for protecting the membrane and protein structures, binding water, securing ions and acting as molecular chaperones (Bray, 1997). Group 3 LEA protein from barley, ABA-inducible HVA1, has been reported by Hong et al. (1988) to sequester ions (e.g., Na+) during cellular dehydration. It has been investigated whether over expressing LEA genes can improve drought tolerance. Higher growth rates and quicker recovery from the stresses demonstrated the enhanced tolerance of the HVA1-transgenic rice and the LEA-transgenic Chinese cabbage to salt stress in potted soil and water deficit conditions during the vegetative stage. By preserving their membrane structure, LEA proteins help stressed cells remain in a suitable cellular environment. Low membrane electrolyte leakage during dehydration stress indicated that transgenic plants had higher cell membrane protection than control plants (Rohila et al., 2002; Babu et al., 2004). The crucial regulatory proteins that are activated during the stress response are known as transcription factors. Besides polymerases, TFs are proteins attaching target gene promoters to control gene expression. According to Joshi et al. (2016), transcription factors are essential for both triggering and regulating the expression of various stress-responsive genes. Different stress responses share certain transcription factors. To fully comprehend abiotic stress tolerance, a thorough examination of transcriptional regulatory systems is necessary. A "Regulon" is a collection of multiple genes activated or inactive by the same transcription factor in response to the same signal. There are two categories for the four main "Regulons" are involved in abiotic stressors, such as drought stress. 1. An ABA-dependent pathway controlled by the AREB/ABF, NAC, and MYB families; 2. An ABA-independent pathway run by DREB TFs carrying the APETALA2 (Flower patterning protein) DNA binding motif (Saibo et al., 2008).

Source: Shankar and Venkateswarlu (2011)

Fig. 1. Classes of genes induced by water-deficit stress.

According to transcriptome studies, numerous primary and secondary stress-responsive genes were found to be expressed in rice exposed to abiotic stresses. Primary stress responsive genes comprise elements of signal transduction pathways such as stress receptors, enzymes that synthesize secondary messengers, MAP kinases, transcription factors and proteins that mitigate stress such as heat shock proteins, late embryogenesis abundant (LEA) proteins, compatible osmolytes and water channels (Aquaporins) (Todaka et al., 2012). In rice and Arabidopsis, both ABA dependent and independent TFs are crucial during drought response (Todaka et al., 2012).

DREB1/CBF regulon: One category of transcription factor not dependent on ABA is DREB1. Three distinct DREB1-type genes-DREB1A, DREB1B and DREB1C-have been identified in Arabidopsis. The core sequence A/GCCGAC is present in the cis-acting sequence that serves as a binding site for these transcription factors. In Arabidopsis, DREB1 TFs are crucial for responding to stresses like cold, high salinity, and drought (Yamaguchi- Shinozaki and Shinozaki, 2005).

DREB2 regulon: Additionally, DREB2 TF functions independently of ABA. Like DREB1, it too has an AP2 domain and is brought on by heat, salinity, and drought. At least four DREB2 homologs have been found in rice, with OsDREB2A and OsDREB2B being induced by heat, salinity, and drought (Matsukura et al., 2010).

AREB regulon: Abiotic stress is a significant time for the ABA hormone. It accumulates during drought-induced water shortages and is an essential signal molecule for stress reactions and tolerance (Fujita et al., 2011).

NAC regulon: NAC transcription factors play a critical in both biotic and abiotic stresses. They make up one of the largest plant-specific TFs. The acronym NAC denotes a set of three distinct genes that share a specific kind of domain known as the NAC domain: Numerous NAC genes have been identified by genome sequencing of crop species, including Vitis, Oryza, Arabidopsis, and citrus (NAM stands for no apical meristem). In rice and Arabidopsis, more than 100 NAC genes have been found (Nakashima et al., 2014; Le et al., 2011). Drought, high salinity and cold stressors activate NAC-type transcription factors, which regulate the expression of multiple stress-responsive genes in Oryza and Arabidopsis.

Calmodulin (CaM) genes induced by drought stress. Calcium binding proteins, a significant class of Ca2+ sensor proteins, are members of the calmodulin (CaM) gene family. These proteins regulate a wide range of target proteins, which play a crucial role in cellular signaling cascades (Reddy et al., 2001; Sanders et al., 2002; White et al., 2003; Ranty et al., 2006). CaM proteins are normally inactive, but when calcium is present, they become active. Elevated concentrations of calcium ions in the cytosol indicate most abiotic stresses. By interacting with calcium sensor proteins like calmodulin, calcium functions as a secondary messenger of the signal transduction pathways and can directly or indirectly activate downstream components. It exists in both the plant and animal kingdoms and is naturally acidic. It is believed to be the primary intracellular Ca2+ receptor in all eukaryotes and is this protein family's most widely distributed member. All eukaryotes share a highly conserved amino acid sequence for CaM proteins (Kawasaki, 1994). Within a single plant species, CaM proteins can exist in multiple isoforms. Four functional EF-hand Ca2+ binding domains are present in plant CaMs.

ASR genes induced by drought stress. ASR (abscisic acid stress ripening) proteins and various polyproteins determine a plant species' stress tolerance. ASR proteins have been repeatedly identified in response to different abiotic stresses, such as drought, salinity, cold or osmotic stress in many plant species (Amitai-Zeigerson et al., 1995; Gilad et al., 1997; Vaidyanathan et al., 1999; Maskin et al., 2008). Notably, each plant species' ability to adapt to water deficit conditions varies greatly and plays a significant role in both productivity and natural geographical distribution. Additionally, ASR is important in controlling the ripening of fruit in strawberries and tomato, where it has been shown that RNAi lines delay fruit ripening and overexpressing ASR lines promote fruit softening and ripening (Jia et al., 2016). For the first time, the tomato's ASR protein was found to be a water/ABA stress-induced protein (Iusem et al., 1993). Additionally, homologs of these proteins have been found in several plant species, such as the lily (Wang et al., 1998), potato (Schneider et al., 1997), melon (Hong et al., 2002), rice (Vaidyanathan et al., 1999), maize (Virlouvet et al., 2011), apricot (Mbeguie-A-Mbeguie et al., 1997), pummelo (Canel et al., 1995), grape (Cakir et al., 2003), strawberry (Chen et al., 2011), and banana (Henry et al., 2011). However, the model plant Arabidopsis thaliana did not recognize these proteins (Carrari et al., 2004). These proteins are hydrophilic and may have a protective function in situations where there is a water deficit (Wang et al., 2005; Konrad and Bar-Zvi 2008). They are members of the low molecular weight charged hydrophylin group. The ASR gene family has different documented members in different plant species, and each member's function varies depending on the host and stress levels. However, the precise function of each member is still unknown. The overexpression of ASR1 genes from wheat and tomatoes in tobacco (Kalifa et al., 2004a; Hu et al., 2013) and the increased tolerance to salt and drought (LLA23) in Arabidopsis (Yang et al., 2005; Hsu et al., 2011) are examples of how the interfamily transfer of these proteins improves tolerance limits. These findings confirm the role of ASR genes in stress response even in systems where they are not endogenously present. Various traits genetically control drought tolerance and is typically inherited quantitatively (Blair et al., 2010).According to Carrari et al. (2004); Philippe et al. (2010), the molecular characterization and sequence comparison revealed that ASR gene members of a single plant species are more closely related than those of other plant species, suggesting that these originated from the late duplication and may have species-specific functions. The majority of ASR proteins consist of two well-conserved regions: the first region, which includes a nuclear localization signal (NLS) at the C-terminal region (Kalifa et al., 2004b), has a stretch of His residues at the N-terminal and contains sequence-specific DNA binding activity (Rom et al., 2006).There are various cellular levels where the ASR proteins are located. ASR proteins confine in the nucleus in various cell types (Kalifa et al., 2004; Wang et al., 2005) and play a regulatory role, possibly as transcription factors or as chaperones (Wang et al., 2005; Frankel et al., 2006; Konrad and Bar-Zvi 2008). The ASR proteins in the cytosol exhibit chaperone-like activity (Konrad and Bar-Zvi 2008). Padaria et al. (2016) have successfully isolated and characterized the ASR gene from Zizypus nummularia, a species of tree that can withstand extreme water stress. When PEG was present, ZnASR1-expressing E. coli demonstrated improved survival under simulated drought stress conditions. ASR4-a recently identified ASR gene from Setaria italica showed a significant increase in expression after treatment with ABA, NaCl and PEG. Additionally, it has been discovered that the ASR4 gene's promoter is bound by the ABA-responsive DRE-binding protein (ARDP). Additionally, A. thaliana's ASR4 gene was altered, and the transgenic lines exhibited improved drought tolerance (Li et al., 2017). Similarly, the rice ASR gene, or ASR5, has also been described. After being isolated from the upland rice variety IRAT109 (Oryza sativa L. ssp. japonica), the ASR5 gene was overexpressed in rice, and transgenic lines with improved drought tolerance were demonstrated by Arabidopsis (Li et al., 2017).

Conclusion

One of the principal abiotic stresses that have a negative impact on agricultural productivity is drought. Significant changes in the global climate in recent years have increased the frequency and severity of droughts. Drought is one of the leading natural causes of severe food shortages in developing nations and is a significant contributing factor to famine and malnutrition. It impacts food security's availability, stability, accessibility and utilization the four pillars. When plants are exposed to environments that restrict water during different developmental stages, various physiological and developmental changes are triggered. There is little understanding of the fundamental biochemical and molecular mechanisms underlying drought stress, transduction, and tolerance. Moreover, genetic engineering is a precious tool for understanding the of drought tolerance mechanism. Because, it can manipulate genes, transcription factors, signaling proteins and genetic regulatory networks that shield plant cells from water deficits. Furthermore, by expanding our understanding of the mechanisms underlying drought, plant breeders have made significant strides towards creating drought-tolerant lines or cultivars for a few key crops. Nevertheless, conventional breeding method is highly labor, time and money-intensive. Because marker-assisted breeding can quickly determine the value of thousands of a crop's genomic regions under stress, it is a more efficient breeding method. By using a technique known as transformation, certain crops with innate mechanisms for surviving droughts can be used as a source of genes for drought tolerance, which can then be used to develop desirable crops.

Future Scope

Present review clearly suggests the significant role of different methods to mitigate the drastic changes in the global climate ecological environment to enhance resistance against both biotic (living) and abiotic (non-living) and organism challenges in the ecological atmosphere. Thus, these methods could be used for developing new resistant genotypes with characteristic properties against various types of stresses in plant systemtomitigate the physical loss of produce the cultural and economic losses in the agricultural and sustainable development.

References

Amitai, Z. H., Scolnik, P. A. and Bar-Zvi, D. (1995). Tomato Asr1 mRNA and protein are transiently expressed following salt stress, osmotic stress and treatment with abscisic acid. Plant Science110(2), 205-213.

Babu, R. C., Zhang, J., Blum, A., Ho, T. H. D., Wu, R. and Nguyen, H. T. (2004). HVA1, a LEA gene from barley confers dehydration tolerance in transgenic rice (Oryza sativa L.) via cell membrane protection. Plant Science166(4), 855-862.

Badawi, G.H., Kawano, N., Yamauchi, Y., Shimada, E., Sasaki, R., Kubo, A. and Tanaka, K. (2004). Over-expression of ascorbate peroxidase in tobacco chloroplasts enhances the tolerance to salt stress and water deficit. Physiologia Plantarum121(2), 231-238.

Bagley, J., Rosenthal, D. M., Ruiz‐Vera, U. M., Siebers, M.H., Kumar, P., Ort, D. R. and Bernacchi, C. J. (2015). The influence of photosynthetic acclimation to rising CO2 and warmer temperatures on leaf and canopy photosynthesis models. Global Biogeochemical Cycles29(2), 194-206.

Bari, R. J. and Jonathan D. G. (2009). "Role of plant hormones in plant defence responses". Plant Molecular Biology, 69 (4), 473–488.

Bartels, D. and Sunkar, R.(2005). Drought and salt tolerance in plants. Critical reviews in plant sciences24(1), 23-58.

Blair, M. W., Galeano, C. H., Tovar, E., Torres, M. C. M., Castrillón, A. V., Beebe, S. E. and Rao, I. M. (2010). Development of a Mesoamerican intra-genepool genetic map for quantitative trait loci detection in a drought tolerant × susceptible common bean (Phaseolus vulgaris L.) cross. Molecular Breeding29(1), 71-88.

Blum, A .(2002). Drought stress and its impact. Plant stress.com. Available from: http://www. plantstress. com/Articles/drought_i/drought_i. htm.

Bray, E. A. (1997). Plant responses to water deficit. Trends in plant science2(2), 48-54.

Burney, J. and Ramanathan, V. (2014). Recent climate and air pollution impacts on Indian agriculture. Proceedings of the National Academy of Sciences111(46), 16319-16324.

Çakir, B., Agasse, A., Gaillard, C., Saumonneau, A., Delrot, S. and Atanassova, R. (2003). A grape ASR protein involved in sugar and abscisic acid signaling. The Plant Cell15(9), 2165-2180.

Canel, C., Bailey-Serres, J. N. and Roose, M. L. (1995). Pummelo fruit transcript homologous to ripening-induced genes. Plant physiology108(3), 1323.

Carrari, F., Fernie, A. R. and Iusem, N. D. (2004). Heard it through the grapevine? ABA and sugar cross-talk: the ASR story. Trends in plant science9(2), 57-59.

Chaves, M. M., Maroco, J. P. and Pereira, J. S. (2003). Understanding plant responses to drought-from genes to the whole plant. Functional plant biology30(3), 239-264.

Checker, V.G., Chhibbar, A.K. and Khurana, P. (2012). Stress-inducible expression of barley Hva1 gene in transgenic mulberry displays enhanced tolerance against drought, salinity and cold stress. Transgenic research21(5), 939-957.

Chen, J. Y., Liu, D. J., Jiang, Y. M., Zhao, M. L., Shan, W., Kuang, J. F. and Lu, W. J. (2011). Molecular characterization of a strawberry FaASR gene in relation to fruit ripening. PloS one6(9), 24649.

Chen, T. H. and Murata, N. (2008). Glycinebetaine: an effective protectant against abiotic stress in plants. Trends in plant science, 13(9), 499-505.

Choudhary, M. and Padaria, J. C. (2015). Transcriptional profiling in pearl millet (Pennisetum glaucum LR Br.) for identification of differentially expressed drought responsive genes. Physiology and Molecular Biology of Plants21(2), 187-196.

Dai, X., Xu, Y., Ma, Q., Xu, W., Wang, T., Xue, Y. and Chong, K. (2007). Overexpression of an R1R2R3 MYB gene, OsMYB3R-2, increases tolerance to freezing, drought, and salt stress in transgenic Arabidopsis. Plant physiology143(4), 1739-1751.

Datta, K., Baisakh, N., Ganguly, M., Krishnan, S., Yamaguchi Shinozaki, K. and Datta, S. K. (2012). Overexpression of Arabidopsis and rice stress genes’ inducible transcription factor confers drought and salinity tolerance to rice. Plant biotechnology journal10(5), 579-586.

De Ronde, J. A., Cress, W. A., Krüger, G. H. J., Strasser, R. J. and Van Staden, J. (2004). Photosynthetic response of transgenic soybean plants, containing an Arabidopsis P5CR gene, during heat and drought stress. Journal of plant physiology161(11), 1211-1224.

De Ronde, J. A., Spreeth, M. H. and Cress, W. A. (2000). Effect of antisense L-1-pyrroline-5-carboxylate reductase transgenic soybean plants subjected to osmotic and drought stress. Plant Growth Regulation32(1), 13-26.

Delauney, A. J. and Verma, D. P. S. (1993). Proline biosynthesis and osmoregulation in plants. The plant journal4(2), 215-223.

Fader, M., Shi, S., Bloh, W. V., Bondeau, A. and Cramer, W. (2016). Mediterranean irrigation under climate change: more efficient irrigation needed to compensate for increases in irrigation water requirements. Hydrology and Earth System Sciences20(2), 953-973.

Flynn, P. (2003). "Biotic vs. Abiotic - Distinguishing Disease Problems from Environmental Stresses". ISU     Entomology.

Frankel, N., Carrari, F., Hasson, E. and Iusem, N. D. (2006). Evolutionary history of the Asr gene family. Gene378, 74-83.

Fujita, Y., Fujita, M., Shinozaki, K. and Yamaguchi-Shinozaki, K. (2011). ABA-mediated transcriptional regulation in response to osmotic stress in plants. Journal of plant research124(4), 509-525.

Gilad, A., Amitai-Zeigerson, H., Bar-Zvi, D. and Scolnik, P. A. (1997). ASR1, a tomato water-stress regulated gene: genomic organization, developmental regulation and DNA-binding activity. In III International Symposium on In Vitro Culture and Horticultural Breeding, 447, 447-454.

He, G. H., Xu, J. Y., Wang, Y. X., Liu, J. M., Li, P. S., Chen, M., Ma, Y. Z. and Xu, Z. S. (2016). Drought-responsive WRKY transcription factor genes TaWRKY1 and TaWRKY33 from wheat confer drought and/or heat resistance in Arabidopsis. BMC plant biology16(1), 116.

Henry, I. M., Carpentier, S. C., Pampurova, S., Van Hoylandt, A., Panis, B., Swennen, R. and Remy, S. (2011). Structure and regulation of the Asr gene family in banana. Planta234(4), 785.

Hirayama, T. and Shinozaki, K. (2010). Research on plant abiotic stress responses in the post-genome era: Past, present and future. The Plant Journal, 61(6), 1041-1052.

Hof, A. R. and Svahlin, A. (2015). The potential effect of climate change on the geographical distribution of insect pest species in the Swedish boreal forest. Scandinavian journal of forest research31(1), 29-39.

Hong, B., Uknes, S. J. and Ho, T. H. D. (1988). Cloning and characterization of a cDNA encoding a mRNA rapidly-induced by ABA in barley aleurone layers. Plant molecular biology, 11(4), 495-506.

Hong, S. H., Kim, I. J., Yang, D. C. and Chung, W. I. (2002). Characterization of an abscisic acid responsive gene homologue from Cucumis meloJournal of experimental botany53(378), 2271-2272.

Hozain, M. D., Abdelmageed, H., Lee, J., Kang, M., Fokar, M., Allen, R. D. and Holaday, A. S. (2012). Expression of AtSAP5 in cotton up-regulates putative stress-responsive genes and improves the tolerance to rapidly developing water deficit and moderate heat stress. Journal of plant physiology169(13), 1261-1270.

Hsu, Y. F., Yu, S. C., Yang, C. Y. and Wang, C. S. (2011). Lily ASR protein-conferred cold and freezing resistance in Arabidopsis. Plant physiology and biochemistry49(9), 937-945.

Hu, W., Huang, C., Deng, X., Zhou, S., Chen, L., Li, Y., Wang, C., Ma, Z., Yuan, Q., Wang, Y. and Cai, R. (2013). TaASR1, a transcription factor gene in wheat, confers drought stress tolerance in transgenic tobacco. Plant, cell & environment36(8), 1449-1464.

Ingram, J. and Bartels, D. (1996). The molecular basis of dehydration tolerance in plants. Annual review of plant biology47(1), 377-403.

Ito, Y., Katsura, K., Maruyama, K., Taji, T., Kobayashi, M., Seki, M., Shinozaki, K. and Yamaguchi-Shinozaki, K. (2006). Functional analysis of rice DREB1/CBF-type transcription factors involved in cold-responsive gene expression in transgenic rice. Plant and Cell Physiology47(1), 141-153.

Iusem, N. D., Bartholomew, D. M., Hitz, W. D. and Scolnik, P. A. (1993). Tomato (Lycopersicon esculentum) transcript induced by water deficit and ripening. Plant Physiology102(4), 1353.

James, D., Tarafdar, A., Biswas, K., Sathyavathi, T. C., Padaria, J. C. and Kumar, P. A. (2015). Development and characterization of a high temperature stress responsive subtractive cDNA library in Pearl Millet Pennisetum glaucum (L.) R. Br. Indian Journal of Experimental Biology, 53, 543-550.

Jia, H., Jiu, S., Zhang, C., Wang, C., Tariq, P., Liu, Z., Wang, B., Cui, L. and Fang, J. (2016). Abscisic acid and sucrose regulate tomato and strawberry fruit ripening through the abscisic acid‐stress‐ripening transcription factor. Plant biotechnology journal, 14(10), 2045-2065.

Joshi, R., Wani, S.H., Singh, B., Bohra, A., Dar, Z. A., Lone, A. A., Pareek, A. and Singla-Pareek, S. L. (2016). Transcription factors and plants response to drought stress: current understanding and future directions. Frontiers in plant science7, 1–15.

Kalifa, Y., Gilad, A., Konrad, Z., Zaccai, M., Scolnik, P. A. and Dudy, B. Z. (2004a). The water-and salt-stress-regulated Asr1 (abscisic acid stress ripening) gene encodes a zinc-dependent DNA-binding protein. Biochemical Journal381(2), 373-378.

Kalifa, Y., Perlson, E., Gilad, A., Konrad, Z., Scolnik, P. A. and Bar-Zvi, D. (2004b). Over-expression of the water and salt stress-regulated Asr1 gene confers an increased salt tolerance. Plant, Cell & Environment27(12), 1459-1468.

Karaba, A., Dixit, S., Greco, R., Aharoni, A., Trijatmiko, K. R., Marsch-Martinez, N., Krishnan, A., Nataraja, K. N., Udayakumar, M. and Pereira, A. (2007). Improvement of water use efficiency in rice by expression of HARDY, an Arabidopsis drought and salt tolerance gene. Proceedings of the National Academy of Sciences104(39), 15270-15275.

Kawasaki, H. (1994). Calcium-binding proteins. 1: EF-hands. Protein profile1, 343-517.

Konrad, Z. and Bar-Zvi, D. (2008). Synergism between the chaperone-like activity of the stress regulated ASR1 protein and the osmolyte glycine-betaine. Planta, 227(6), 1213-1219.

Krasensky, J. and Jonak, C. (2012). Drought, salt, and temperature stress-induced metabolic rearrangements and regulatory networks. Journal of experimental botany63(4), 1593-1608.

Le, D. T., Nishiyama, R. I. E., Watanabe, Y., Mochida, K., Yamaguchi-Shinozaki, K., Shinozaki, K. and Tran, L.S. P. (2011). Genome-wide survey and expression analysis of the plant-specific NAC transcription factor family in soybean during development and dehydration stress. DNA research18(4), 263-276.

Lee, S. B., Kim, H., Kim, R. J. and Suh, M. C. (2014). Overexpression of Arabidopsis MYB96 confers drought resistance in Camelina sativa via cuticular wax accumulation. Plant cell reports33(9), 1535-1546.

Leila, R. (2007). Response of Tunisian autochthonous pearl millet (Pennisetum glaucum (L.) R. Br.) to drought stress induced by polyethylene glycol (PEG) 6000. African journal of biotechnology6(9), 1102-1105.

Levitt, J. (1972). Response of plants to environmental stresses. Water, radiation, Salt and other stressesAcademic Press: New York. 698.

Li, J., Li, Y., Yin, Z., Jiang, J., Zhang, M., Guo, X., Ye, Z., Zhao, Y., Xiong, H., Zhang, Z. and Shao, Y. (2017). OsASR5 enhances drought tolerance through a stomatal closure pathway associated with ABA and H2O2 signalling in rice. Plant biotechnology journal15(2), 183-196.

Luo, L. J. (2010). Breeding for water-saving and drought-resistance rice (WDR) in China. Journal of experimental botany61(13), 3509-3517.

Lv, S., Yang, A., Zhang, K., Wang, L. and Zhang, J. (2007). Increase of glycinebetaine synthesis improves drought tolerance in cotton. Molecular Breeding, 20(3), 233-248.

Ma, T., Li, M., Zhao, A., Xu, X., Liu, G. and Cheng, L. (2014). LcWRKY5: an unknown function gene from sheepgrass improves drought tolerance in transgenic Arabidopsis. Plant cell reports, 33(9), 1507-1518.

Manavalan, L. P., Guttikonda, S. K., Phan Tran, L. S. and Nguyen, H. T. (2009). Physiological and molecular approaches to improve drought resistance in soybean. Plant and Cell Physiology50(7), 1260-1276.

Maskin, L., Maldonado, S. and Iusem, N. D. (2008). Tomato leaf spatial expression of stress-induced Asr genes. Molecular biology reports35(4), 501-505.

Matsukura, S., Mizoi, J., Yoshida, T., Todaka, D., Ito, Y., Maruyama, K., Shinozaki, K. and Yamaguchi-Shinozaki, K. (2010). Comprehensive analysis of rice DREB2-type genes that encode transcription factors involved in the expression of abiotic stress-responsive genes. Molecular Genetics and Genomics283(2), 185-196.

Mazel, A., Leshem, Y., Tiwari, B. S. and Levine, A. (2004). Induction of salt and osmotic stress tolerance by overexpression of an intracellular vesicle trafficking protein AtRab7 (AtRabG3e). Plant physiology, 134(1), 118-128.

Mbeguie-A-Mbeguie, D., Gomez, R. M. and Fils-Lycaon, B. (1997). Molecular cloning and nucleotide sequence of an abscisic acid-, stress-, ripening-induced (ASR)-like protein from apricot fruit (accession no. U82760). Gene expression during fruit ripening (PGR97–161). Plant Physiol115, 1288.

Mishra, R. N., Reddy, P. S., Nair, S., Markandeya, G., Reddy, A. R., Sopory, S. K. and Reddy, M. K. (2007). Isolation and characterization of expressed sequence tags (ESTs) from subtracted cDNA libraries of Pennisetum glaucum seedlings. Plant molecular biology64(6), 713-732.

Mitra, J. (2001). Genetics and genetic improvement of drought resistance in crop plants. Current science, 758-763.

Monte, L. M. (1986). Breeding plants for drought resistance-the problem and its relevance.Drought resistance in plants-Physiological and genetic aspects. Amalfi. October 19th to 23rd, 1-11.

Mooney, H. A., Pearcy, R. W. and Ehleringer, J. (1987). How plants cope: Plant physiological ecology, 37(1), 18-20.

Nakashima, K., Yamaguchi-Shinozaki, K. and Shinozaki, K.(2014). The transcriptional regulatory network in the drought response and its crosstalk in abiotic stress responses including drought, cold, and heat. Frontiers in plant science5(170), 1-7.

Nelson, D. E., Repetti, P. P., Adams, T. R., Creelman, R.A., Wu, J., Warner, D. C., Anstrom, D. C., Bensen, R. J., Castiglioni, P. P., Donnarummo, M. G. and Hinchey, B. S. (2007). Plant nuclear factor Y (NF-Y) B subunits confer drought tolerance and lead to improved corn yields on water-limited acres. Proceedings of the National Academy of Sciences104(42), 16450-16455.

Oh, S. J., Song, S. I., Kim, Y.S., Jang, H. J., Kim, S. Y., Kim, M., Kim, Y. K., Nahm, B. H. and Kim, J. K. (2005). Arabidopsis CBF3/DREB1A and ABF3 in transgenic rice increased tolerance to abiotic stress without stunting growth. Plant physiology138(1), 341-351.

Ozturk, A. (1998). The effect of drought on the growth and yield of winter wheat. Turkish Journal of Agriculture and Forestry23(5), 531-540.

Padaria, J. C., Yadav, R., Tarafdar, A., Lone, S. A., Kumar, K. and Sivalingam, P. N. (2016). Molecular cloning and characterization of drought stress responsive abscisic acid-stress-ripening (Asr. Molecular biology reports43(8), 849-859.

Passioura, J. B. (1997). Drought and drought tolerance. In Drought tolerance in higher plants: genetical, physiological and molecular biological analysis, p1-5. Springer Netherlands.

Pennisi, E. (2008). Plant Genetics; The Blue revolution, drop by drop, gene by gene. News Focus Science230, 171-173.

Peterson, R. K. D., Higley, L. G., eds. (2001). Biotic Stress and Yield Loss. CRC Press. 

Philippe, R., Courtois, B., McNally, K. L., Mournet, P., El-Malki, R., Le Paslier, M. C., Fabre, D., Billot, C., Brunel, D., Glaszmann, J. C. and This, D. (2010). Structure, allelic diversity and selection of Asr genes, candidate for drought tolerance, in Oryza sativa L. and wild relatives. Theoretical and applied genetics121(4), 769-787.

Pilon-Smits, E. A., Ebskamp, M. J., Paul, M. J., Jeuken, M. J., Weisbeek, P. J. and Smeekens, S. C. (1995). Improved performance of transgenic fructan-accumulating tobacco under drought stress. Plant physiology107(1), 125-130.

Price, A. H., Cairns, J. E., Horton, P., Jones, H. G. and Griffiths, H. (2002). Linking drought-resistance mechanisms to drought avoidance in upland rice using a QTL approach: progress and new opportunities to integrate stomatal and mesophyll responses. Journal of Experimental Botany53(371), 989-1004.

Quan, R., Shang, M., Zhang, H., Zhao, Y. and Zhang, J. (2004). Engineering of enhanced glycine betaine synthesis improves drought tolerance in maize. Plant Biotechnology Journal2(6), 477-486. 

Ramegowda, V. and Senthil-Kumar, M. (2015). The interactive effects of simultaneous biotic and abiotic stresses on plants: mechanistic understanding from drought and pathogen combination. Journal of plant physiology176, 47-54.

Ranty, B., Aldon, D. and Galaud, J. P. (2006). Plant calmodulins and calmodulin-related proteins: multifaceted relays to decode calcium signals. Plant signaling & behavior1(3), 96-104.

Rao, T. S. R. B., Naresh, J.V., Reddy, P. S., Reddy, M. K. and Mallikarjuna, G. (2017). Expression of Pennisetum glaucum Eukaryotic Translational Initiation Factor 4A (PgeIF4A) Confers Improved Drought, Salinity, and Oxidative Stress Tolerance in Groundnut. Frontiers in plant science, 8.

Reddy, A. R., Chaitanya, K. V. and Vivekanandan, M.(2004). Drought-induced responses of photosynthesis and antioxidant metabolism in higher plants. Journal of plant physiology161(11), 1189-1202.

Reddy, A. S. (2001). Calcium: silver bullet in signaling. Plant Science160(3), 381-404.

Reis, R. R., da Cunha, B. A. D. B., Martins, P. K., Martins, M. T. B., Alekcevetch, J. C., Chalfun-Júnior, A., Andrade, A. C., Ribeiro, A. P., Qin, F., Mizoi, J. and Yamaguchi-Shinozaki, K. (2014). Induced over-expression of AtDREB2A CA improves drought tolerance in sugarcane. Plant Science221, 59-68.

Rohila, J. S., Jain, R. K. and Wu, R. (2002). Genetic improvement of Basmati rice for salt and drought tolerance by regulated expression of a barley Hva1 cDNA. Plant Science163(3), 525-532.

Rom, S., Gilad, A., Kalifa, Y., Konrad, Z., Karpasas, M.M., Goldgur, Y. and Bar-Zvi, D. (2006). Mapping the DNA-and zinc-binding domains of ASR1 (abscisic acid stress ripening), an abiotic-stress regulated plant specific protein. Biochimie, 88(6), 621-628.

Saibo, N. J., Lourenço, T. and Oliveira, M. M. (2008). Transcription factors and regulation of photosynthetic and related metabolism under environmental stresses. Annals of botany103(4), 609-623.

Sanders, D., Pelloux, J., Brownlee, C. and Harper, J. F. (2002). Calcium at the crossroads of signaling. The Plant Cell14(suppl 1), S401-S417.

Schneider, A., Salamini, F. and Gebhardt, C. (1997). Expression patterns and promoter activity of the cold-regulated gene ci21A of potato. Plant physiology113(2), 335-345.

Scott, J. W., Volin, R. B., Bryan, H. H. and Olson, S. M. (1986). Use of hybrids to develop heat tolerant tomato cultivars. In Proceedings of the annual meeting of the Florida State Horticulture Society (USA).

Seki, M., Narusaka, M., Ishida, J., Nanjo, T., Fujita, M., Oono, Y., Kamiya, A., Nakajima, M., Enju, A., Sakurai, T. and Satou, M. (2002). Monitoring the expression profiles of 7000 Arabidopsis genes under drought, cold and high-salinity stresses using a full-length cDNA microarray. The Plant Journal, 31(3), 279-292.

Shanker, A. and Venkateswarlu, B. (2011). Abiotic Stress Response in Plants-Physiological, Biochemical and Genetic Perspectives. In Tech publisher.

Sharma-Natu, P. and Ghildiyal, M. C. (2005). Potential targets for improving photosynthesis and crop yield. Current Science, 1918-1928.

Shinozaki, K. and Yamaguchi-Shinozaki, K. (2000). Molecular responses to dehydration and low temperature: differences and cross-talk between two stress signaling pathways. Current opinion in plant biology3(3), 217-223.

Shou, H., Bordallo, P. and Wang, K. (2004). Expression of the Nicotiana protein kinase (NPK1) enhanced drought tolerance in transgenic maize. Journal of experimental botany55(399), 1013-1019.

Spoel, S. H. and Dong, X., (2012). How do plants achieve immunity? Defence without specialized immune cells. Nature Reviews Immunology, 12(2), 89–100.

Sun, W., Bernard, C., Van De Cotte, B., Van Montagu, M. and Verbruggen, N. (2001). At-HSP17. 6A, encoding a small heat-shock protein in Arabidopsis, can enhance osmotolerance upon overexpression. The Plant Journal27(5), 407-415.

Taiz, L., Zeiger, E., Moller, I. M. and Murphy, A. (2015). Plant Physiology and Development. USA: Sinauer Associations, Inc. p. 706. 

Tang, Y., Liu, M., Gao, S., Zhang, Z., Zhao, X., Zhao, C., Zhang, F. and Chen, X. (2012). Molecular characterization of novel TaNAC genes in wheat and overexpression of TaNAC2a confers drought tolerance in tobacco. Physiologiaplantarum144(3), 210-224.

Tester, M. and Bacic, A. (2005). Abiotic stress tolerance in grasses. From model plants to crop plants. Plant Physiology137(3), 791-793.

Todaka, D., Nakashima, K., Shinozaki, K. and Yamaguchi-Shinozaki, K. (2012). Toward understanding transcriptional regulatory networks in abiotic stress responses and tolerance in rice. Rice5(1), 6.

Tripathy, J. N., Zhang, J., Robin, S., Nguyen, T. T. and Nguyen, H. T. (2000). QTLs for cell-membrane stability mapped in rice (Oryza sativa L.) under drought stress. TAG Theoretical and Applied Genetics, 100(8), 1197-1202.

Tsuda, K. and Katagiri, F. (2010). "Comparing signaling mechanisms engaged in pattern-triggered and effector-triggered immunity". Current Opinion in Plant Biology, 13 (4) 459–465.

Twidwell, E. K. (2002). Effects of drought stress on soybean production. http://agbiopubs.Sdstate.edu/articles/ Ex 8034 pdf.

Vaidyanathan, R., Kuruvilla, S. and Thomas, G. (1999). Characterization and expression pattern of an abscisic acid and osmotic stress responsive gene from rice. Plant Science140(1), 21-30.

Valente, M. A. S., Faria, J. A., Soares-Ramos, J. R., Reis, P. A., Pinheiro, G. L., Piovesan, N. D., Morais, A. T., Menezes, C. C., Cano, M. A., Fietto, L. G. and Loureiro, M. E. (2009). The ER luminal binding protein (BiP) mediates an increase in drought tolerance in soybean and delays drought-induced leaf senescence in soybean and tobacco. Journal of experimental botany60(2), 533-546.

Virlouvet, L., Jacquemot, M. P., Gerentes, D., Corti, H., Bouton, S., Gilard, F., Valot, B., Trouverie, J., Tcherkez, G., Falque, M. and Damerval, C. (2011). The ZmASR1 protein influences branched-chain amino acid biosynthesis and maintains kernel yield in maize under water-limited conditions. Plant physiology157(2), 917-936.

Wang, C. S., Liau, Y. E., Huang, J.C., Wu, T.D., Su, C. C. and Lin, C. H. (1998). Characterization of a desiccation-related protein in lily pollen during development and stress. Plant and cell physiology39(12), 1307-1314.

Wang, H. J., Hsu, C. M., Jauh, G. Y. and Wang, C. S. (2005). A lily pollen ASR protein localizes to both cytoplasm and nuclei requiring a nuclear localization signal. Physiologiaplantarum123(3), 314-320.

Wani, H., Shabir, Singh, N. B., Haribhushan, A. and Mir, J. I. (2013). Compatible solute engineering in plants for abiotic stress tolerance-role of glycine betaine. Current genomics, 14(3), 157-165.

White, P. J. and Broadley, M. R. (2003). Calcium in plants. Annals of botany, 92(4), 487-511.

Yamaguchi-Shinozaki, K. and Shinozaki, K. (2005). Organization of cis-acting regulatory elements in osmotic and cold-stress-responsive promoters. Trends in plant science10(2), 88-94.

Yang Han, Y., xiu Li, A., Li, F., rong Zhao, M. and Wang, W. (2012). Characterization of a wheat (Triticum aestivum L.) expansin gene, TaEXPB23, involved in the abiotic stress response and phytohormone regulation. Plant Physiology and Biochemistry54, 49-58.

Yang, C. Y., Chen, Y. C., Jauh, G. Y. and Wang, C. S. (2005). A lily ASR protein involves abscisic acid signaling and confers drought and salt resistance in Arabidopsis. Plant physiology139(2), 836-846.

Yang, M., Yang, Q., Fu, T. and Zhou, Y. (2011). Overexpression of the Brassica napus BnLAS gene in Arabidopsis affects plant development and increases drought tolerance. Plant cell reports30(3), 373-388.

Yoshida, T., Fujita, Y., Sayama, H., Kidokoro, S., Maruyama, K., Mizoi, J., Shinozaki, K. and Yamaguchi-Shinozaki, K. (2010). AREB1, AREB2, and ABF3 are master transcription factors that cooperatively regulate ABRE-dependent ABA signaling involved in drought stress tolerance and require ABA for full activation. The Plant Journal, 61(4), 672-685.

Yue, B., Xue, W., Xiong, L., Yu, X., Luo, L., Cui, K., Jin, D., Xing, Y. and Zhang, Q. (2006). Genetic basis of drought resistance at reproductive stage in rice: separation of drought tolerance from drought avoidance. Genetics172(2), 1213-1228.

Zandalinas, S.I., Sales, C., Beltrán, J., Gómez-Cadenas, A. and Arbona, V. (2017). Activation of secondary metabolism in citrus plants is associated to sensitivity to combined drought and high temperatures. Frontiers in plant science7, 1-17.

Zhang, G., Liu, X., Quan, Z., Cheng, S., Xu, X., Pan, S., Xie, M., Zeng, P., Yue, Z., Wang, W. and Tao, Y. (2012). Genome sequence of foxtail millet (Setaria italica) provides insights into grass evolution and biofuel potential. Nature biotechnology, 30(6), 549-554.

Zhang, H., Mao, X., Wang, C. and Jing, R. (2010). Overexpression of a common wheat gene TaSnRK2. 8 enhances tolerance to drought, salt and low temperature in Arabidopsis. PloS one, 5(12), 16041.

Zhu, J. K. (2002). Salt and drought stress signal transduction in plants. Annual review of plant biology53(1), 247-273.

Zhu, J. K., Hasegawa, P. M., Bressan, R. A. and Bohnert, H. J. (1998). Molecular aspects of osmotic stress in plants. Critical Reviews in Plant Sciences16(3), 253-277.

Zörb, C., Schmitt, S., Neeb, A., Karl, S., Linder, M. and Schubert, S. (2004). The biochemical reaction of maize (Zea mays L.) to salt stress is characterized by a mitigation of symptoms and not by a specific adaptation. Plant Science167(1), 91-100.

How to cite this article

Rajendra Prasad Meena, Deen Dayal Bairwa, Vikas Sharma, Sanjiv Kumar, Satyesh Raj Anand, Harkesh Kumar Balai,Vijay Daneva and Ashish Kumar Sharma  (2024). Identification of Different Stress Responsive Genes Induced in Crops for Climate Changes: A Review. Biological Forum – An International Journal, 16(5): 06-17.