Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions

*Kemal Yuce and Ahmet Ismail Ozkan*

### **Abstract**

Zinc finger (ZnF) proteins are the largest transcription factors family. They constitute of nine sub-groups including Cys2His2, Cys3His, Cys3HisCys4, Cys2HisCys5, Cys4HisCys3, Cys2HisCys, Cys4, Cys6 and Cys8. ZnFs perform tasks of recognizing DNA, packaging RNA, transcriptional activity, regulating apoptosis, folding and collecting proteins, and binding lipids. One of the largest sub-groups of these proteins is ZF-Cys2His2, containing SIZ1, ZAT, ZAT7, ZFP1, ZFP252, DST, ZFP1, SIZF3, ZFP179, ZjZFN1, SICZFP1, and ZF-Cys2His2 proteins are found in plants tissues and fulfill important tasks in their defense to struggle with biotic and abiotic stresses (i.e., salt, drought, cold, oxidative). The aim of this chapter is to reveal importance of ZF-Cys2His2 proteins against various stress conditions.

**Keywords:** Cys2His2 Zinc finger proteins, plant stress physiology, salinity stress, cold stress, oxidative stress

### **1. Introduction**

Cys2His2 zinc finger (ZF-Cys2His2) proteins have been found in a number of plants including *Arabidopsis*, cotton, rice and wheat. The ZF-Cys2His2 is built by two Cys and two His residues. This structure provides a conservative motif together with Zn2+. The core has an -helix and an anti-parallel double-stranded -sheet so that the ZF-Cys2His2 proteins have stable and relatively independent protein domains. ZF-Cys2His2 proteins form a relatively wide family of transcriptional factors in plants. Current studies have revealed that ZF-Cys2His2 proteins operate as important transcriptional regulators in plant responses to a large broad of stress conditions (like drought, excessive light, extreme temperatures, oxidative stress, salinity, and silique shattering) [1]. Over-expression of some ZF-Cys2His2 genes has led to an increased tolerance to various stresses and activation of some stressrelated genes [2]. Abscisic acid (ABA) is an important phytohormone involved in regulating stress responses and plant growth and development. In addition, ABA is involved in many important processes of plants such as stomatal closure, leaf senescence, cuticle wax accumulation, bud dormancy, osmotic regulation, seed germination and growth inhibition. Since the ABA regulates downstream responses to biotic and abiotic environmental changes through both transcriptional and post-transcription mechanisms, the responses of ABA and ZF-Cys2His2 proteins to various stresses are also mentioned here [3].

### *Plant Stress Physiology*

The world population has exceeded 7.5 billion. To supply the nutritional needs of this population, it is important to know the proteins and genes related to the response of plants to stress conditions. In this context, to increase the durability and yield of plants, transgenic plant researches are carried out. And the most commonly focused proteins in these researches are ZF-Cys2His2 proteins. For this reason, in this chapter, the changes occurring in ZF-Cys2His2 proteins with transgenic methods and various stress conditions and what these changes bring to the plant have been discussed (**Table 1**).


**191**

*Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions*

One of the stress factors is salt. Plants are affected with development and yield from salt stress. Transgenic plant studies in combating salt stress have become one of the most important issues of our century and ZnF proteins attract a lot of attention in this context. Salt application has induced expression of AtSIZ1. The germination energy, index and rate, cotyledon growth rate and root length were found to be importantly higher than wild-type in lines where AtSIZ1 was over-expressed under various stress applications at the germination stage. However, these indicators decreased significantly in AtSIZ1 mutants. Higher proline, potassium and soluble sugar, lower sodium, malondialdehyde, sodium/potassium ratios were observed in the lines where over-expression occurred in the mature seedling stage compared to the wild-type. Stress-related marker genes such as AtGSTU5, AtP5CS1, COR15A, RD29A, RD29B and SOS1 have been found to be over-expressed in lines with an excessive expression than those of wild-type and mutant under salt application. Due to the results AtSIZ1 functions in maintaining both ionic homeostasis and osmotic balance to improve salt tolerance in *Arabidopsis* [4]. It has also been shown that GmZAT4 plays a significant role in both polyethylene glycol and sodium chloride stresses tolerance and ABA responses in both *A. thaliana* and soybean. Over-expression of GmZAT4 increased the tolerance of *A. thaliana* to 20% polyethylene glycol and 150 mM sodium chloride and increased the germination rate after 1 or 2 μM ABA administration [5]. The EAR motif for ZAT7 has a significant role of defense response to salt stress in *Arabidopsis*. Suppressing growth and being more tolerant to salt stress have observed in transgenic *Arabidopsis* plants expressing ZF-Cys2His2 protein ZAT7. Mutation or deletion of ZAT7's EAR motif did not affect growth suppression, but eliminated salt tolerance. These results showed that ZAT7's EAR motif is important in increasing salt stress tolerance of the transgenic plants. On the other hand, the EAR motif appears not to be involved in suppressing transgenic plants growth. Other analyzes of ZAT7 using RNAi lines suggested that ZAT7 functions as a suppressor of the defense response's suppressor [6]. Semiquantitative RT-PCR studies have revealed that AhZFP1 expression is stimulated by salt stress in the peanut root, stem and leaves [7]. Excessive expression of ZFP252 in rice increased the amount of free proline, soluble sugar, stress defense genes expression and improved the plant's response to both salt and drought stress. This result showed that ZFP252 plays a significant role in both salt and drought stresses of rice plant. And it is also useful for genetically modified plants to increase both salt and drought stresses tolerance [8]. ZmZF1 transcripts are strongly stimulated by salt stress. It is also stimulated by drought stress and ABA. Over-expression of ZmZF1 under cauliflower mosaic virus 35S promoter control increased both salt and drought tolerance in young seedling phase of transgenic *Arabidopsis* [9]. Stomata control the absorption of carbon-dioxide and improve water use efficiency. Thus, it plays an important role in abiotic stress tolerance. H2O2, stimulates stomatal closure, is an important signaling molecule. Another ZFP, DST is directly organizing genes related to H2O2 homeostasis. Therefore, the stomatal closure is regulated negatively. The loss of DST function decreases stomatal density and increases stomatal closure. As a result, increased salt and drought tolerance appears in the rice plant. These findings provide a different perspective on the abiotic stress tolerance mechanism associated with stomata and also a significant genetic engineering approach in developing abiotic stress tolerance [10]. GhZFP1 over-expression in transgenic tobacco plant increased salt stress tolerance and *Rhizoctonia solani* resistance. This indicates that the plant can act as a significant regulator in responding to biotic and abiotic stress [11]. Ascorbic acid supports stress tolerance by breaking down reactive oxygen species (ROS). ROS degradation capacity of ascorbic acid is increased

*DOI: http://dx.doi.org/10.5772/intechopen.92590*

**2. Salinity stress and Cys2His2 zinc finger**

### **Table 1.**

*C2H2 zinc finger proteins related with plant stress.*

### **2. Salinity stress and Cys2His2 zinc finger**

*Plant Stress Physiology*

have been discussed (**Table 1**).

The world population has exceeded 7.5 billion. To supply the nutritional needs of this population, it is important to know the proteins and genes related to the response of plants to stress conditions. In this context, to increase the durability and yield of plants, transgenic plant researches are carried out. And the most commonly focused proteins in these researches are ZF-Cys2His2 proteins. For this reason, in this chapter, the changes occurring in ZF-Cys2His2 proteins with transgenic methods and various stress conditions and what these changes bring to the plant

**The kind of C2H2 zinc finger proteins The kind of stress References**

Salt, cold and drought stress [20]

Oxidative stress [27]

[28]

AtSIZ1 Salt stress [4] GmZAT4 Salt stress [5] ZAT7 Salt stress [6] AhZFP1 Salt stress [7] ZFP252 Salt stress [8] ZmZF1 Salt stress [9] DST Salt stress [10] GhZFP1 Salt stress [11] SIZF3 Salt stress [12, 13] ZFP179 Salt stress [14] ZjZFN1 Salt and cold stress [15] TaZNF Salt stress [16] SICZFP1 Salt and cold stress [17] TaDi19A Salt osmotic and cold stress [18] AtDi19-3 Salt and drought stress [19]

GsZFP1 Cold and drought stress [21] ZFP245 Cold and drought stress [22] GbZF1 Cold stress [23] PeSTZ1 Cold and oxidative stress [24] SCOF-1 Cold stress [25] OsCTZFP8 Cold stress [26]

ZFP36 Oxidative stress [29] GsZFP1 Cold stress [21] ZAT18 Oxidative and drought stress [30] OsMSR15 Drought stress [31] VTA2 Oxidative and fungal stress [32] MtSTOP pH and aluminum stress [33] ART1 Aluminum stress [34]

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**Table 1.**

*C2H2 zinc finger proteins related with plant stress.*

AZF2 STZ

ZAT12 ZAT7

One of the stress factors is salt. Plants are affected with development and yield from salt stress. Transgenic plant studies in combating salt stress have become one of the most important issues of our century and ZnF proteins attract a lot of attention in this context. Salt application has induced expression of AtSIZ1. The germination energy, index and rate, cotyledon growth rate and root length were found to be importantly higher than wild-type in lines where AtSIZ1 was over-expressed under various stress applications at the germination stage. However, these indicators decreased significantly in AtSIZ1 mutants. Higher proline, potassium and soluble sugar, lower sodium, malondialdehyde, sodium/potassium ratios were observed in the lines where over-expression occurred in the mature seedling stage compared to the wild-type. Stress-related marker genes such as AtGSTU5, AtP5CS1, COR15A, RD29A, RD29B and SOS1 have been found to be over-expressed in lines with an excessive expression than those of wild-type and mutant under salt application. Due to the results AtSIZ1 functions in maintaining both ionic homeostasis and osmotic balance to improve salt tolerance in *Arabidopsis* [4]. It has also been shown that GmZAT4 plays a significant role in both polyethylene glycol and sodium chloride stresses tolerance and ABA responses in both *A. thaliana* and soybean. Over-expression of GmZAT4 increased the tolerance of *A. thaliana* to 20% polyethylene glycol and 150 mM sodium chloride and increased the germination rate after 1 or 2 μM ABA administration [5]. The EAR motif for ZAT7 has a significant role of defense response to salt stress in *Arabidopsis*. Suppressing growth and being more tolerant to salt stress have observed in transgenic *Arabidopsis* plants expressing ZF-Cys2His2 protein ZAT7. Mutation or deletion of ZAT7's EAR motif did not affect growth suppression, but eliminated salt tolerance. These results showed that ZAT7's EAR motif is important in increasing salt stress tolerance of the transgenic plants. On the other hand, the EAR motif appears not to be involved in suppressing transgenic plants growth. Other analyzes of ZAT7 using RNAi lines suggested that ZAT7 functions as a suppressor of the defense response's suppressor [6]. Semiquantitative RT-PCR studies have revealed that AhZFP1 expression is stimulated by salt stress in the peanut root, stem and leaves [7]. Excessive expression of ZFP252 in rice increased the amount of free proline, soluble sugar, stress defense genes expression and improved the plant's response to both salt and drought stress. This result showed that ZFP252 plays a significant role in both salt and drought stresses of rice plant. And it is also useful for genetically modified plants to increase both salt and drought stresses tolerance [8]. ZmZF1 transcripts are strongly stimulated by salt stress. It is also stimulated by drought stress and ABA. Over-expression of ZmZF1 under cauliflower mosaic virus 35S promoter control increased both salt and drought tolerance in young seedling phase of transgenic *Arabidopsis* [9]. Stomata control the absorption of carbon-dioxide and improve water use efficiency. Thus, it plays an important role in abiotic stress tolerance. H2O2, stimulates stomatal closure, is an important signaling molecule. Another ZFP, DST is directly organizing genes related to H2O2 homeostasis. Therefore, the stomatal closure is regulated negatively. The loss of DST function decreases stomatal density and increases stomatal closure. As a result, increased salt and drought tolerance appears in the rice plant. These findings provide a different perspective on the abiotic stress tolerance mechanism associated with stomata and also a significant genetic engineering approach in developing abiotic stress tolerance [10]. GhZFP1 over-expression in transgenic tobacco plant increased salt stress tolerance and *Rhizoctonia solani* resistance. This indicates that the plant can act as a significant regulator in responding to biotic and abiotic stress [11]. Ascorbic acid supports stress tolerance by breaking down reactive oxygen species (ROS). ROS degradation capacity of ascorbic acid is increased

in plants that over-expressed SIZF3. This has increased the salt tolerance of plants. Findings in the studies showed that SIZF3 simultaneously supports both the collection of ascorbic acid and improves the plant salt tolerance response [12]. ZFP3 expression level was found to be highly suppressed by mannitol, sodium chloride and sucrose. The mutant *Arabidopsis* exhibited a reduced tolerance condition in terms of ZFP3. Excessive expression of ZFP3 reduced stress-related genes (such as AtP5CS1, KIN1, RD22 and RD29B) expression. These results showed that ZFP3 is important in osmotic and salt stress response [13]. Over-expression of ZFP179 increased salt tolerance in rice plant. And the transgenic seedlings showed hypersensitive activity to exogenic ABA. Increased amount of soluble sugars and free proline under salt stress have been observed in transgenic plants. In ZFP179 transgenic rice plant the oxidative stress tolerance, ability to break down ROS and expression level of stress-related genes such as OsDREB2A, OsLea3, OsP5CS and OsProT increased under salt stress. These findings showed that ZFP179 plays an important role in salt stress in plants and is useful for developing transgenic plants that are highly tolerant to salt stress [14]. Expression of ZjZFN1 was found to be increased by ABA, cold and salt. ZjZFN1 expression improved seed germination, plant adaptation to salt stress by growth under salt stress and increased green cotyledons in *Arabidopsis thaliana*. Physiological and transcriptional analyzes suggest that ZjZFN1 can affect the collection of ROS and regulate the transcription of genes associated with salt response. ZjZFN1 over-expressing plants RNA sequence analysis revealed that ZjZFN1 could function as a transcriptional activator in the regulation of stress response pathways such as alpha-linoleic acid metabolism, phenyl alanine metabolism and phenylpropanoid biosynthesis pathways. The results showed that ZjZFN1 plays an important role in the formation of salt stress response in plants [15]. In another study, it has been proven that TaZnF, a ZF-Cys2His2 protein, significantly improves salt tolerance of transgenic *Arabidopsis*. Various physiological indexes of transgenic plants showed improvement under salt stress compared to the control group. The noninvasive micro-test (NMT) technique showed the excretion of Na is significantly accomplished by the root end of the transgenic *Arabidopsis*. TaZnF is mainly found in the nucleus and has demonstrated transcriptional activity [16].

### **3. Cold stress and C2H2 zinc finger proteins**

GsZFP1 was found to be stimulated by ABA (100 μM), cold (4°C) and salt (200 mM sodium chloride) in leaves and by ABA (100 μM), cold (4°C) and drought (30% PEG 6000) in root. It was found that over-expression of GsZFP1 in transgenic *Arabidopsis* resulted in higher tolerance in cold and drought stress and a reduced rate of water loss. Over-expression of GsZFP1 was found to increase the expression of CBF1, CBF2, CBF3, COR47, NCED3 and RD29A stress response genes in response to cold stress, and increased expression of NCED3, P5CS, RAB18, RD22 and RD29A in response to drought stress [17]. It was revealed by the semi-quantitative RT-PCR experiment that ZFP245 was strongly stimulated after 6 hours of cold and drought stress and then decreased to normal level. ZFP245 did not occur by the application of high salt or ABA [18]. High degree of aggregation of GmZF1 mRNAs stimulated by exogenic ABA suggested that GmZF1 was involved in the ABA-dependent signal transduction pathway. GmZF1 over-expression increased expression of the cold-induced cor6.6 gene by recognizing the likely protein DNA binding site. This suggests that GbZF1 of soybean can improve *Arabidopsis* cold stress tolerance by regulating cold regulatory genes expression in transgenic *Arabidopsis* [19]. PeSTZ1 increases freezing tolerance through organizing the breakdown of ROS by directly regulating PeAPX. PeSTZ1 is preferably expressed in young roots. This ZFP has

**193**

*Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions*

been upregulated in cold applications. PeSTZ1 functions as a transcriptional activator to increase cold tolerance. PeSTZ1 upregulation decreases malondialdehyde and ROS aggregation by activating antioxidant systems. This is thought to be achieved through direct regulation of PeAPX2 expression [20]. SCOF-1 transcription is particularly stimulated by ABA and low temperature, but not neither with dehydration nor high salt. SCOF-1 over-expression stimulated the expression of genes regulated by cold and improved the cold tolerance of tobacco plants and non-acclimated transgenic *Arabidopsis.* SCOF-1 can function as a positive regulator of COR gene expression regulated by ABRE through protein-protein interaction and thus can improve the cold tolerance of plants [21]. SICZFP1 is strongly stimulated by cold stress, dehydration and salt stress, but not by ABA. SICZFP1 over-expression in transgenic rice and *Arabidopsis* plants increased expression of cold-response-related genes. This suggests that SICZFP1 plays an important role in the response of plants to cold stress [22]. TaDi19A has been found to be expressed in both leaves and roots of wheat seedlings grown under stress-free conditions, but is significantly upregulated in salt, osmotic and cold stress conditions, or in hormone applications such as stress-related ethylene and ABA. Heterologous over-expression of TaDi19A in *Arabidopsis thaliana* increased salinity, mannitol and ABA stress sensitivity during the germination of plants. Root elongation in these transgenic lines showed less salinity stress tolerance and less ethephon sensitivity. The expression of ABA signal path genes such as ABA1, ABF3, ERD15, RAB18 and SOS2 (SOS pathway) have changed in transgenic plants [23]. Transgenic rice expressing excessive OsCTZFP8 exhibited cold-tolerant phenotypes with significantly higher pollen fertility and seed placement rates than non-transgenic control plants. Yield of lines expressing OsCTZFP8 per plant was significantly higher (p < 0.01) than non-transgenic control plants under cold application. This result shows that OsCTZFP8 is a C2H2 zinc finger transcription factor that plays an important role in cold tolerance in rice [24].

*DOI: http://dx.doi.org/10.5772/intechopen.92590*

**4. Oxidative stress and C2H2 zinc finger proteins**

Cytosolic Apx1, ascorbate peroxidase 1, is an important H2O2-removing enzyme

in plants. Both WRKY transcription factor (WRKY25) and two ZnF proteins (ZAT12 and ZAT7) expressions have increased in Apx1 gene suppressed plants that grown under the controlled conditions. When cells were exposed to oxidative stress, heat shock, and injury, the WRKY25, ZAT7 and ZAT12 expressions increased together. However, light and osmotic stresses did not increase them. Transgenic plants expressing ZAT7 and ZAT12 were able to tolerate oxidative stress. WRKY25, ZAT7 or ZAT12 expression in transgenic plants did not increase Apx1 expression under controlled conditions. Plants without ZAT12 could not increase Apx1, WRKY25 and ZAT7 expressions in response to H2O2, so that plants without ZAT12 have become more sensitive to H2O2 applications than normal plants. It has been revealed that ZAT12 is an important component of oxidative stress signal transmission in *Arabidopsis* and needs Apx1, WRKY25 and ZAT7 during oxidative stress [25]. Transcription profiles of plants that are exposed to H2O2 stress, expressing excessive ZAT12 and normal have revealed that the basic expression of ZAT12 in *Arabidopsis* results in increased expression in oxidative and mild stress transcripts. Thus, ZAT12 has been shown to play a key role in reactive oxygen and abiotic stress signal in *Arabidopsis* [26]. ABA application stimulated the increase of both ascorbate peroxidase and superoxide dismutase activities and OsMPK1, OsMPK5 and ZFP182 expressions in rice plant leaves. It has been noted that ABA-induced antioxidant defense needs ZFP182. And the ZFP182 expression is regulated by rice MAPKs in ABA signaling [27]. ABA-activated mitogen active protein kinases

*Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions DOI: http://dx.doi.org/10.5772/intechopen.92590*

been upregulated in cold applications. PeSTZ1 functions as a transcriptional activator to increase cold tolerance. PeSTZ1 upregulation decreases malondialdehyde and ROS aggregation by activating antioxidant systems. This is thought to be achieved through direct regulation of PeAPX2 expression [20]. SCOF-1 transcription is particularly stimulated by ABA and low temperature, but not neither with dehydration nor high salt. SCOF-1 over-expression stimulated the expression of genes regulated by cold and improved the cold tolerance of tobacco plants and non-acclimated transgenic *Arabidopsis.* SCOF-1 can function as a positive regulator of COR gene expression regulated by ABRE through protein-protein interaction and thus can improve the cold tolerance of plants [21]. SICZFP1 is strongly stimulated by cold stress, dehydration and salt stress, but not by ABA. SICZFP1 over-expression in transgenic rice and *Arabidopsis* plants increased expression of cold-response-related genes. This suggests that SICZFP1 plays an important role in the response of plants to cold stress [22]. TaDi19A has been found to be expressed in both leaves and roots of wheat seedlings grown under stress-free conditions, but is significantly upregulated in salt, osmotic and cold stress conditions, or in hormone applications such as stress-related ethylene and ABA. Heterologous over-expression of TaDi19A in *Arabidopsis thaliana* increased salinity, mannitol and ABA stress sensitivity during the germination of plants. Root elongation in these transgenic lines showed less salinity stress tolerance and less ethephon sensitivity. The expression of ABA signal path genes such as ABA1, ABF3, ERD15, RAB18 and SOS2 (SOS pathway) have changed in transgenic plants [23]. Transgenic rice expressing excessive OsCTZFP8 exhibited cold-tolerant phenotypes with significantly higher pollen fertility and seed placement rates than non-transgenic control plants. Yield of lines expressing OsCTZFP8 per plant was significantly higher (p < 0.01) than non-transgenic control plants under cold application. This result shows that OsCTZFP8 is a C2H2 zinc finger transcription factor that plays an important role in cold tolerance in rice [24].

### **4. Oxidative stress and C2H2 zinc finger proteins**

Cytosolic Apx1, ascorbate peroxidase 1, is an important H2O2-removing enzyme in plants. Both WRKY transcription factor (WRKY25) and two ZnF proteins (ZAT12 and ZAT7) expressions have increased in Apx1 gene suppressed plants that grown under the controlled conditions. When cells were exposed to oxidative stress, heat shock, and injury, the WRKY25, ZAT7 and ZAT12 expressions increased together. However, light and osmotic stresses did not increase them. Transgenic plants expressing ZAT7 and ZAT12 were able to tolerate oxidative stress. WRKY25, ZAT7 or ZAT12 expression in transgenic plants did not increase Apx1 expression under controlled conditions. Plants without ZAT12 could not increase Apx1, WRKY25 and ZAT7 expressions in response to H2O2, so that plants without ZAT12 have become more sensitive to H2O2 applications than normal plants. It has been revealed that ZAT12 is an important component of oxidative stress signal transmission in *Arabidopsis* and needs Apx1, WRKY25 and ZAT7 during oxidative stress [25]. Transcription profiles of plants that are exposed to H2O2 stress, expressing excessive ZAT12 and normal have revealed that the basic expression of ZAT12 in *Arabidopsis* results in increased expression in oxidative and mild stress transcripts. Thus, ZAT12 has been shown to play a key role in reactive oxygen and abiotic stress signal in *Arabidopsis* [26]. ABA application stimulated the increase of both ascorbate peroxidase and superoxide dismutase activities and OsMPK1, OsMPK5 and ZFP182 expressions in rice plant leaves. It has been noted that ABA-induced antioxidant defense needs ZFP182. And the ZFP182 expression is regulated by rice MAPKs in ABA signaling [27]. ABA-activated mitogen active protein kinases

*Plant Stress Physiology*

in plants that over-expressed SIZF3. This has increased the salt tolerance of plants. Findings in the studies showed that SIZF3 simultaneously supports both the collection of ascorbic acid and improves the plant salt tolerance response [12]. ZFP3 expression level was found to be highly suppressed by mannitol, sodium chloride and sucrose. The mutant *Arabidopsis* exhibited a reduced tolerance condition in terms of ZFP3. Excessive expression of ZFP3 reduced stress-related genes (such as AtP5CS1, KIN1, RD22 and RD29B) expression. These results showed that ZFP3 is important in osmotic and salt stress response [13]. Over-expression of ZFP179 increased salt tolerance in rice plant. And the transgenic seedlings showed hypersensitive activity to exogenic ABA. Increased amount of soluble sugars and free proline under salt stress have been observed in transgenic plants. In ZFP179 transgenic rice plant the oxidative stress tolerance, ability to break down ROS and expression level of stress-related genes such as OsDREB2A, OsLea3, OsP5CS and OsProT increased under salt stress. These findings showed that ZFP179 plays an important role in salt stress in plants and is useful for developing transgenic plants that are highly tolerant to salt stress [14]. Expression of ZjZFN1 was found to be increased by ABA, cold and salt. ZjZFN1 expression improved seed germination, plant adaptation to salt stress by growth under salt stress and increased green cotyledons in *Arabidopsis thaliana*. Physiological and transcriptional analyzes suggest that ZjZFN1 can affect the collection of ROS and regulate the transcription of genes associated with salt response. ZjZFN1 over-expressing plants RNA sequence analysis revealed that ZjZFN1 could function as a transcriptional activator in the regulation of stress response pathways such as alpha-linoleic acid metabolism, phenyl alanine metabolism and phenylpropanoid biosynthesis pathways. The results showed that ZjZFN1 plays an important role in the formation of salt stress response in plants [15]. In another study, it has been proven that TaZnF, a ZF-Cys2His2 protein, significantly improves salt tolerance of transgenic *Arabidopsis*. Various physiological indexes of transgenic plants showed improvement under salt stress compared to the control group. The noninvasive micro-test (NMT) technique showed the excretion of Na is significantly accomplished by the root end of the transgenic *Arabidopsis*. TaZnF is mainly found in the nucleus and has demonstrated transcriptional activity [16].

**3. Cold stress and C2H2 zinc finger proteins**

GsZFP1 was found to be stimulated by ABA (100 μM), cold (4°C) and salt (200 mM sodium chloride) in leaves and by ABA (100 μM), cold (4°C) and drought (30% PEG 6000) in root. It was found that over-expression of GsZFP1 in transgenic *Arabidopsis* resulted in higher tolerance in cold and drought stress and a reduced rate of water loss. Over-expression of GsZFP1 was found to increase the expression of CBF1, CBF2, CBF3, COR47, NCED3 and RD29A stress response genes in response to cold stress, and increased expression of NCED3, P5CS, RAB18, RD22 and RD29A in response to drought stress [17]. It was revealed by the semi-quantitative RT-PCR experiment that ZFP245 was strongly stimulated after 6 hours of cold and drought stress and then decreased to normal level. ZFP245 did not occur by the application of high salt or ABA [18]. High degree of aggregation of GmZF1 mRNAs stimulated by exogenic ABA suggested that GmZF1 was involved in the ABA-dependent signal transduction pathway. GmZF1 over-expression increased expression of the cold-induced cor6.6 gene by recognizing the likely protein DNA binding site. This suggests that GbZF1 of soybean can improve *Arabidopsis* cold stress tolerance by regulating cold regulatory genes expression in transgenic *Arabidopsis* [19]. PeSTZ1 increases freezing tolerance through organizing the breakdown of ROS by directly regulating PeAPX. PeSTZ1 is preferably expressed in young roots. This ZFP has

**192**

(MAPKs) and ABA-induced H2O2 production have been shown to regulate ZFP36 expression in ABA signaling, on the other hand, NADPH oxidase genes expression regulated by ZFP36, H2O2 production and the expression of OsMPK genes in ABA signaling. It has been noted that rice plants requires to ZFP36 for tolerance of both water stress and oxidative stresses, for ABA-induced antioxidant defense and for cross-interaction between NADPH oxidase, H2O2 and MAPK in the ABA signal [28].

### **5. Drought stress and C2H2 zinc finger proteins**

Over-expression of ZAT18 has increased drought tolerance in *Arabidopsis* while ZAT18 mutation reduced plant tolerance to drought stress. It was found that ZAT18 is expressed in stems and herbal rosette leaves. Sub-cellularly, it has been noted that ZAT18 protein is mainly localized in the nucleus. Overexpressed ZAT18 plants exhibited higher leaf water content, lower ROS content, higher antioxidant enzyme activity compared to wild-type after drought application. RNA sequencing analysis showed that the 423 and 561 genes were transcriptionally regulated by the ZAT18 transgene before and after drought application, respectively. Path enrichment analysis showed that hormone metabolism, stress and signal were over represented in ZAT18 overexpression lines. Various stress-sensitive genes (including COR47, ERD7, LEA6 and RAS1) and hormone signal transduction-related genes (including JAZ7 and PYL5) have been identified as target genes of ZAT18. Taken together, ZAT18 acts as a positive regulator and plays an important role in the plant's response to drought stress [29]. Di19 family, protein 19 induced by drought, is a ZF-Cys2His2 protein. AtDi19-3 is a transcription activator. In plants, AtDi19-3 is significantly stimulated by ABA, mannitol and sodium chloride. Mutation of AtDi19-3 increased tolerance of plants against ABA, drought and high salinity. Excessive expression of AtDi19-3 caused drought, salt and ABA sensitivity. Seed germination and cotyledon greening rates were higher in the Atdi19-3 mutant under the condition of applying sodium chloride, mannitol or ABA. But this data in transgenic plants, in which AtDi19-3 was over-expressed, was lower than in normal plants. Although the roots of AtDi19-3 mutant seedings were longer, the roots of AtDi19-3 over-expressing transgenic seedlings were shorter than in normal plants. Although the chlorophyll and proline contents in the AtDi19-3 mutant were higher, these data of AtDi19-3 over-expressed seedlings were lower than that of normal plants. While the AtDi19-3 mutant tolerates more droughts, transgenic plants that over-express AtDi19-3 showed more susceptibility to drought than normal plants. In addition, ABA related genes expression signaling pathway varied in the Atdi19-3 mutant and in over-expressing plants of AtDi19-3 [30]. AZF2 and STZ expression has been shown to be strongly stimulated by ABA, cold and high salt stresses, and dehydration. Transgenic *Arabidopsis*, which over-expresses STZ, has become tolerant to growth delay and drought stress. In that study, it was suggested that STZ and AZF2 act as transcriptional suppressors on increased stress tolerance during the growth delay [31]. The expression of OsMSR15 in *Arabidopsis* provides it with drought resistance. Hypersensitive activity to exogenic ABA in this transgenic plant growth during sprouting and seed germination. Transgenic plants also showed less electrolyte leakage, high levels of free proline and increased expressions of a number of stress sensitive genes (such as LEA3, DREB1A, P5CS1 and RD29A) under drought stress. The results showed that OsMSR15 is a significant regulator that plays a role against plant drought stress [32]. In another study, over-expression of TaZFP34 has been shown to have a negative effect on yield performance and mimic the negative effect of drought stress on wheat productivity [33]. STF-2 is also a ZF-Cys2His2 protein.

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*Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions*

And its transgenically over-expression increased significantly drought tolerance in transgenic tobacco. The results showed that STF-2 plays a significant role in the

VTA2 contains the ZF-Cys2His2 proteins. It controls H2O2 detoxification and host plant root infection. And it is an important regulator of fungal pathogenesis [35]. StZFP2 is a Q-type C2H2 zinc finger transcription factor induced by injury and invasion. Previous studies show that Q-type C2H2 TFs are involved in responding to stress and may be protective against drought, salinity, or pathogenic infections when overexpressed. The increase in StPIN2, a classic marker for insect defense in potatoes, was consistent with decreases in larval weight gain [36]. MtSTOP is

toxicity. MtSTOP is expressed in root, stem, nodule and other tissues. MtSTOP is upregulated with acidic pH and Al3+ stress or a combination of both. Growth or morphology in mtSTOP mutants did not change under normal conditions; however, mutant seedlings are characterized by significantly reduced root elongation and are sensitive to low pH (pH 4.3) and Al3+ stress. Compared to its control, more Al accumulated in the mutant roots and citric acid secreted from the mutant roots was significantly lower in both normal and Al stress conditions. This indicates that MtSTOP hair roots synthesize more citric and malic acids [37]. Regarding aluminum, Rice (*Oryza sativa*) is one of the aluminum tolerant species among small grain cereals. This type of aluminum tolerance occurs with many genes involved in the detoxification of Al at the cellular level. These findings emerged with ART1. ART1 is a C2H2 type zinc finger transcription factor and regulates the expression of 31

This work was supported by the Selcuk University and Dicle University.

and Al3+

also a C2H2 zinc finger protein and regulates Medicago's response to H+

*DOI: http://dx.doi.org/10.5772/intechopen.92590*

response of soybean's to drought stress [34].

genes as down-stream [38].

The authors declare no conflict of interest.

**Conflict of interest**

**Financial support**

**6. Other stress and C2H2 zinc finger proteins**

And its transgenically over-expression increased significantly drought tolerance in transgenic tobacco. The results showed that STF-2 plays a significant role in the response of soybean's to drought stress [34].

## **6. Other stress and C2H2 zinc finger proteins**

VTA2 contains the ZF-Cys2His2 proteins. It controls H2O2 detoxification and host plant root infection. And it is an important regulator of fungal pathogenesis [35]. StZFP2 is a Q-type C2H2 zinc finger transcription factor induced by injury and invasion. Previous studies show that Q-type C2H2 TFs are involved in responding to stress and may be protective against drought, salinity, or pathogenic infections when overexpressed. The increase in StPIN2, a classic marker for insect defense in potatoes, was consistent with decreases in larval weight gain [36]. MtSTOP is also a C2H2 zinc finger protein and regulates Medicago's response to H+ and Al3+ toxicity. MtSTOP is expressed in root, stem, nodule and other tissues. MtSTOP is upregulated with acidic pH and Al3+ stress or a combination of both. Growth or morphology in mtSTOP mutants did not change under normal conditions; however, mutant seedlings are characterized by significantly reduced root elongation and are sensitive to low pH (pH 4.3) and Al3+ stress. Compared to its control, more Al accumulated in the mutant roots and citric acid secreted from the mutant roots was significantly lower in both normal and Al stress conditions. This indicates that MtSTOP hair roots synthesize more citric and malic acids [37]. Regarding aluminum, Rice (*Oryza sativa*) is one of the aluminum tolerant species among small grain cereals. This type of aluminum tolerance occurs with many genes involved in the detoxification of Al at the cellular level. These findings emerged with ART1. ART1 is a C2H2 type zinc finger transcription factor and regulates the expression of 31 genes as down-stream [38].

### **Conflict of interest**

*Plant Stress Physiology*

(MAPKs) and ABA-induced H2O2 production have been shown to regulate ZFP36 expression in ABA signaling, on the other hand, NADPH oxidase genes expression regulated by ZFP36, H2O2 production and the expression of OsMPK genes in ABA signaling. It has been noted that rice plants requires to ZFP36 for tolerance of both water stress and oxidative stresses, for ABA-induced antioxidant defense and for cross-interaction between NADPH oxidase, H2O2 and MAPK in the ABA signal [28].

Over-expression of ZAT18 has increased drought tolerance in *Arabidopsis* while ZAT18 mutation reduced plant tolerance to drought stress. It was found that ZAT18 is expressed in stems and herbal rosette leaves. Sub-cellularly, it has been noted that ZAT18 protein is mainly localized in the nucleus. Overexpressed ZAT18 plants exhibited higher leaf water content, lower ROS content, higher antioxidant enzyme activity compared to wild-type after drought application. RNA sequencing analysis showed that the 423 and 561 genes were transcriptionally regulated by the ZAT18 transgene before and after drought application, respectively. Path enrichment analysis showed that hormone metabolism, stress and signal were over represented in ZAT18 overexpression lines. Various stress-sensitive genes (including COR47, ERD7, LEA6 and RAS1) and hormone signal transduction-related genes (including JAZ7 and PYL5) have been identified as target genes of ZAT18. Taken together, ZAT18 acts as a positive regulator and plays an important role in the plant's response to drought stress [29]. Di19 family, protein 19 induced by drought, is a ZF-Cys2His2 protein. AtDi19-3 is a transcription activator. In plants, AtDi19-3 is significantly stimulated by ABA, mannitol and sodium chloride. Mutation of AtDi19-3 increased tolerance of plants against ABA, drought and high salinity. Excessive expression of AtDi19-3 caused drought, salt and ABA sensitivity. Seed germination and cotyledon greening rates were higher in the Atdi19-3 mutant under the condition of applying sodium chloride, mannitol or ABA. But this data in transgenic plants, in which AtDi19-3 was over-expressed, was lower than in normal plants. Although the roots of AtDi19-3 mutant seedings were longer, the roots of AtDi19-3 over-expressing transgenic seedlings were shorter than in normal plants. Although the chlorophyll and proline contents in the AtDi19-3 mutant were higher, these data of AtDi19-3 over-expressed seedlings were lower than that of normal plants. While the AtDi19-3 mutant tolerates more droughts, transgenic plants that over-express AtDi19-3 showed more susceptibility to drought than normal plants. In addition, ABA related genes expression signaling pathway varied in the Atdi19-3 mutant and in over-expressing plants of AtDi19-3 [30]. AZF2 and STZ expression has been shown to be strongly stimulated by ABA, cold and high salt stresses, and dehydration. Transgenic *Arabidopsis*, which over-expresses STZ, has become tolerant to growth delay and drought stress. In that study, it was suggested that STZ and AZF2 act as transcriptional suppressors on increased stress tolerance during the growth delay [31]. The expression of OsMSR15 in *Arabidopsis* provides it with drought resistance. Hypersensitive activity to exogenic ABA in this transgenic plant growth during sprouting and seed germination. Transgenic plants also showed less electrolyte leakage, high levels of free proline and increased expressions of a number of stress sensitive genes (such as LEA3, DREB1A, P5CS1 and RD29A) under drought stress. The results showed that OsMSR15 is a significant regulator that plays a role against plant drought stress [32]. In another study, over-expression of TaZFP34 has been shown to have a negative effect on yield performance and mimic the negative effect of drought stress on wheat productivity [33]. STF-2 is also a ZF-Cys2His2 protein.

**5. Drought stress and C2H2 zinc finger proteins**

**194**

The authors declare no conflict of interest.

## **Financial support**

This work was supported by the Selcuk University and Dicle University.

*Plant Stress Physiology*

### **Author details**

Kemal Yuce1 \* and Ahmet Ismail Ozkan<sup>2</sup>

1 Faculty of Medicine, Department of Physiology, Selcuk University, Konya, Turkey

2 Graduate School of Natural and Applied Sciences, Department of Biology, Dicle University, Diyarbakir, Turkey

\*Address all correspondence to: yuce@selcuk.edu.tr

© 2020 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

**197**

*Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions*

tolerance in rice (*Oryza sativa* L.). FEBS

Letters. 2008;**582**(7):1037-1043

[10] Huang XY, Chao DY, Gao JP, Zhu MZ, Shi M, Lin HX. A previously unknown zinc finger protein, DST, regulates drought and salt tolerance in rice via stomatal aperture control. Genes & Development. 2009;**23**(15):1805-1817

[11] Guo Y-H, Yu Y-P, Wang D, Wu C-A, Yang G-D, Huang J-G, et al. GhZFP1, a novel CCCH-type zinc finger protein from cotton, enhances salt stress tolerance and fungal disease resistance in transgenic tobacco by interacting with GZIRD21A and GZIPR5. New Phytologist. 2009;**183**(1):62-75

[12] Li Y, Chu Z, Luo J, Zhou Y, Cai Y, Lu Y, et al. The C2H2 zinc-finger protein SlZF3 regulates AsA synthesis and salt tolerance by interacting with CSN5B. Plant Biotechnology Journal.

[13] Wu K, Zhang A, Liu D, Hua C, Yan A, Liu B, et al. The *Arabidopsis* gene zinc finger protein 3(ZFP3) is involved in salt stress and

osmotic stress response. PLOS One.

[14] Sun SJ, Guo SQ, Yang X, Bao YM, Tang HJ, Sun H, et al. Functional analysis of a novel Cys2/His2-type zinc finger protein involved in salt tolerance in rice. Journal of Experimental Botany.

[15] Teng K, Tan P, Guo W, Yue Y, Fan X, Wu J. Heterologous expression of a novel *Zoysia japonica* C2H2 zinc finger

2018;**16**(6):1201-1213

2016;**11**(12):e0168367

2010;**61**(10):2807-2818

[9] Huai J, Zheng J, Wang G. Overexpression of a new Cys2/His2 zinc finger protein ZmZF1 from maize confers salt and drought tolerance in transgenic *Arabidopsis*. Plant Cell, Tissue and Organ Culture (PCTOC).

2009;**99**(2):117-124

*DOI: http://dx.doi.org/10.5772/intechopen.92590*

[1] Wang K, Ding Y, Cai C, Chen Z, Zhu C. The role of C2H2 zinc finger proteins in plant responses to abiotic stresses. Physiologia Plantarum.

[2] Kiełbowicz-Matuk A. Involvement of plant C2H2-type zinc finger

transcription factors in stress responses. Plant Science. 2012;**185-186**:78-85

[3] Chen K, Li GJ, Bressan RA, Song CP, Zhu JK, Zhao Y. Abscisic acid dynamics, signaling, and functions in plants. Journal of Integrative Plant Biology.

[4] Han G, Yuan F, Guo J, Zhang Y, Sui N, Wang B. AtSIZ1 improves salt tolerance by maintaining ionic homeostasis and osmotic balance in *Arabidopsis*. Plant Science.

[5] Sun Z, Liu R, Guo B, Huang K, Wang L, Han Y, et al. Ectopic expression of GmZAT4, a putative C2H2-type zinc finger protein, enhances PEG and NaCl stress tolerances in *Arabidopsis thaliana*.

[6] Ciftci-Yilmaz S, Morsy MR, Song L, Coutu A, Krizek BA, Lewis MW, et al. The EAR-motif of the Cys2/His2-type zinc finger protein Zat7 plays a key role in the defense response of *Arabidopsis* to salinity stress. Journal of Biological Chemistry. 2007;**282**(12):9260-9268

[7] Pan L, Yang Q, Chi X, Chen M, He Y, Yu S. AhZEP1, a cDNA encoding C2H2 type zinc finger protein, induced by salt stress in peanut (*Arachis hypogaea* L.). In: 2010 4th International Conference on Bioinformatics and Biomedical

[8] Xu D-Q, Huang J, Guo S-Q, Yang X, Bao Y-M, Tang H-J, et al. Overexpression of a TFIIIA-type zinc finger protein geneZFP252enhances drought and salt

Engineering. 2010. pp. 1-7

**References**

2018;**165**(4):690-700

2020;**62**(1):25-54

2019;**285**:55-67

3 Biotech. 2019;**9**(5)

*Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions DOI: http://dx.doi.org/10.5772/intechopen.92590*

### **References**

*Plant Stress Physiology*

**196**

**Author details**

University, Diyarbakir, Turkey

\* and Ahmet Ismail Ozkan<sup>2</sup>

\*Address all correspondence to: yuce@selcuk.edu.tr

provided the original work is properly cited.

1 Faculty of Medicine, Department of Physiology, Selcuk University, Konya, Turkey

2 Graduate School of Natural and Applied Sciences, Department of Biology, Dicle

© 2020 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

Kemal Yuce1

[1] Wang K, Ding Y, Cai C, Chen Z, Zhu C. The role of C2H2 zinc finger proteins in plant responses to abiotic stresses. Physiologia Plantarum. 2018;**165**(4):690-700

[2] Kiełbowicz-Matuk A. Involvement of plant C2H2-type zinc finger transcription factors in stress responses. Plant Science. 2012;**185-186**:78-85

[3] Chen K, Li GJ, Bressan RA, Song CP, Zhu JK, Zhao Y. Abscisic acid dynamics, signaling, and functions in plants. Journal of Integrative Plant Biology. 2020;**62**(1):25-54

[4] Han G, Yuan F, Guo J, Zhang Y, Sui N, Wang B. AtSIZ1 improves salt tolerance by maintaining ionic homeostasis and osmotic balance in *Arabidopsis*. Plant Science. 2019;**285**:55-67

[5] Sun Z, Liu R, Guo B, Huang K, Wang L, Han Y, et al. Ectopic expression of GmZAT4, a putative C2H2-type zinc finger protein, enhances PEG and NaCl stress tolerances in *Arabidopsis thaliana*. 3 Biotech. 2019;**9**(5)

[6] Ciftci-Yilmaz S, Morsy MR, Song L, Coutu A, Krizek BA, Lewis MW, et al. The EAR-motif of the Cys2/His2-type zinc finger protein Zat7 plays a key role in the defense response of *Arabidopsis* to salinity stress. Journal of Biological Chemistry. 2007;**282**(12):9260-9268

[7] Pan L, Yang Q, Chi X, Chen M, He Y, Yu S. AhZEP1, a cDNA encoding C2H2 type zinc finger protein, induced by salt stress in peanut (*Arachis hypogaea* L.). In: 2010 4th International Conference on Bioinformatics and Biomedical Engineering. 2010. pp. 1-7

[8] Xu D-Q, Huang J, Guo S-Q, Yang X, Bao Y-M, Tang H-J, et al. Overexpression of a TFIIIA-type zinc finger protein geneZFP252enhances drought and salt

tolerance in rice (*Oryza sativa* L.). FEBS Letters. 2008;**582**(7):1037-1043

[9] Huai J, Zheng J, Wang G. Overexpression of a new Cys2/His2 zinc finger protein ZmZF1 from maize confers salt and drought tolerance in transgenic *Arabidopsis*. Plant Cell, Tissue and Organ Culture (PCTOC). 2009;**99**(2):117-124

[10] Huang XY, Chao DY, Gao JP, Zhu MZ, Shi M, Lin HX. A previously unknown zinc finger protein, DST, regulates drought and salt tolerance in rice via stomatal aperture control. Genes & Development. 2009;**23**(15):1805-1817

[11] Guo Y-H, Yu Y-P, Wang D, Wu C-A, Yang G-D, Huang J-G, et al. GhZFP1, a novel CCCH-type zinc finger protein from cotton, enhances salt stress tolerance and fungal disease resistance in transgenic tobacco by interacting with GZIRD21A and GZIPR5. New Phytologist. 2009;**183**(1):62-75

[12] Li Y, Chu Z, Luo J, Zhou Y, Cai Y, Lu Y, et al. The C2H2 zinc-finger protein SlZF3 regulates AsA synthesis and salt tolerance by interacting with CSN5B. Plant Biotechnology Journal. 2018;**16**(6):1201-1213

[13] Wu K, Zhang A, Liu D, Hua C, Yan A, Liu B, et al. The *Arabidopsis* gene zinc finger protein 3(ZFP3) is involved in salt stress and osmotic stress response. PLOS One. 2016;**11**(12):e0168367

[14] Sun SJ, Guo SQ, Yang X, Bao YM, Tang HJ, Sun H, et al. Functional analysis of a novel Cys2/His2-type zinc finger protein involved in salt tolerance in rice. Journal of Experimental Botany. 2010;**61**(10):2807-2818

[15] Teng K, Tan P, Guo W, Yue Y, Fan X, Wu J. Heterologous expression of a novel *Zoysia japonica* C2H2 zinc finger

gene, ZjZFN1, improved salt tolerance in *Arabidopsis*. Frontiers in Plant Science. 2018;**9**

[16] Ma X, Liang W, Gu P, Huang Z. Salt tolerance function of the novel C2H2-type zinc finger protein TaZNF in wheat. Plant Physiology and Biochemistry. 2016;**106**:129-140

[17] Luo X, Bai X, Zhu D, Li Y, Ji W, Cai H, et al. GsZFP1, a new Cys2/ His2-type zinc-finger protein, is a positive regulator of plant tolerance to cold and drought stress. Planta. 2011;**235**(6):1141-1155

[18] Huang J, Wang JF, Wang QH, Zhang HS. Identification of a rice zinc finger protein whose expression is transiently induced by drought, cold but not by salinity and abscisic acid. DNA Sequence. 2009;**16**(2):130-136

[19] Chen F, Yu G-H, Jiang L-L, Ma X-F, Xu Z-S, Liu M-M, et al. A soybean C2H2-type zinc finger gene GmZF1 enhanced cold tolerance in transgenic *Arabidopsis*. PLOS One. 2014;**9**(10):e109399

[20] He F, Li HG, Wang JJ, Su Y, Wang HL, Feng CH, et al. Pe STZ 1, a C2H2-type zinc finger transcription factor from *Populus euphratica*, enhances freezing tolerance through modulation of ROS scavenging by directly regulating Pe APX 2. Plant Biotechnology Journal. 2019;**17**(11):2169-2183

[21] Kim JC, Lee SH, Cheong YH, Yoo C-M, Lee SI, Chun HJ, et al. A novel cold-inducible zinc finger protein from soybean, SCOF-1, enhances cold tolerance in transgenic plants. The Plant Journal. 2001;**25**(3):247-259

[22] Zhang X, Guo X, Lei C, Cheng Z, Lin Q, Wang J, et al. Overexpression of SlCZFP1, a novel TFIIIA-type zinc finger protein from tomato, confers enhanced cold tolerance in transgenic *Arabidopsis* and rice. Plant Molecular Biology Reporter. 2010;**29**(1):185-196

[23] Li S, Xu C, Yang Y, Xia G. Functional analysis ofTaDi19A, a saltresponsive gene in wheat. Plant, Cell & Environment. 2010

[24] Jin Y-M, Piao R, Yan Y-F, Chen M, Wang L, He H, et al. Overexpression of a new zinc finger protein transcription factors CTZFP8 improves cold tolerance in rice. International Journal of Genomics. 2018;**2018**:1-13

[25] Rizhsky L, Davletova S, Liang H, Mittler R. The zinc finger protein Zat12 is required for cytosolic ascorbate peroxidase 1 expression during oxidative stress in *Arabidopsis*. Journal of Biological Chemistry. 2004;**279**(12):11736-11743

[26] Davletova S, Schlauch K, Coutu J, Mittler R. The zinc-finger protein Zat12 plays a central role in reactive oxygen and abiotic stress signaling in *Arabidopsis*. Plant Physiology. 2005;**139**(2):847-856

[27] Zhang H, Ni L, Liu Y, Wang Y, Zhang A, Tan M, et al. The C2H2-type zinc finger protein ZFP182 is involved in abscisic acid-induced antioxidant defense in rice. Journal of Integrative Plant Biology. 2012;**54**(7):500-510

[28] Zhang H, Liu Y, Wen F, Yao D, Wang L, Guo J, et al. A novel rice C2H2 type zinc finger protein, ZFP36, is a key player involved in abscisic acid-induced antioxidant defence and oxidative stress tolerance in rice. Journal of Experimental Botany. 2014;**65**(20):5795-5809

[29] Yin M, Wang Y, Zhang L, Li J, Quan W, Yang L, et al. The *Arabidopsis* Cys2/His2 zinc finger transcription factor ZAT18 is a positive regulator of plant tolerance to drought stress. Journal of Experimental Botany. 2017;**68**(11):2991-3005

**199**

*Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions*

[37] Wang J, Hou Q, Li P, Yang L, Sun X, Benedito VA, et al. Diverse functions of multidrug and toxin extrusion (MATE) transporters in citric acid efflux and metal homeostasis in *Medicago truncatula*. The Plant Journal.

[38] Tsutsui T, Yamaji N, Feng MJ. Identification of a cis-acting element of ART1, a C2H2-type zinc-finger transcription factor for aluminum tolerance in rice. Plant Physiology.

2017;**90**(1):79-95

2011;**156**(2):925-931

*DOI: http://dx.doi.org/10.5772/intechopen.92590*

[31] Sakamoto H, Maruyama K, Sakuma Y, Meshi T, Iwabuchi M, Shinozaki K, et al. *Arabidopsis* Cys2/His2-type zinc-finger proteins function as transcription repressors under drought, cold, and high-salinity stress conditions. Plant Physiology. 2004;**136**(1):2734-2746

[32] Zhang X, Zhang B, Li MJ, Yin XM, Huang LF, Cui YC, et al. OsMSR15 encoding a rice C2H2-type zinc finger protein confers enhanced drought tolerance in transgenic *Arabidopsis*. Journal of Plant Biology.

[34] Song B, Zhang Y, Li Y, Fu Y, Wang P. Novel drought-inducible Cys2/His2-type zinc finger protein STF-2 from soybean (*Glycine max*) enhances drought tolerance in transgenic plants. Pakistan

2016;**59**(3):271-281

[33] Chang H, Chen D, Kam J, Richardson T, Drenth J, Guo X, et al. Abiotic stress upregulated TaZFP34 represses the expression of type-B response regulator and SHY2 genes and enhances root to shoot ratio in wheat. Plant Science. 2016;**252**:88-102

Journal of Botany. 2019;**51**(3)

[35] Tran V-T, Braus-Stromeyer SA, Kusch H, Reusche M, Kaever A,

[36] Lawrence SD, Novak NG. Overexpression of StZFP2 in *Solanum tuberosum* L. var. Kennebec (potato) inhibits growth of Tobacco Hornworm larvae (THW, *Manduca sexta* L.). Plant Signaling & Behavior. 2018;**13**(7):e1489668

Kühn A, et al. Verticillium transcription activator of adhesion Vta2 suppresses microsclerotia formation and is required for systemic infection of plant roots. New Phytologist. 2014;**202**(2):565-581

[30] Qin LX, Li Y, Li DD, Xu WL, Zheng Y, Li XB. *Arabidopsis* droughtinduced protein Di19-3 participates in plant response to drought and high salinity stresses. Plant Molecular Biology. 2014;**86**(6):609-625

*Cys2His2 Zinc Finger Proteins Boost Survival Ability of Plants against Stress Conditions DOI: http://dx.doi.org/10.5772/intechopen.92590*

[30] Qin LX, Li Y, Li DD, Xu WL, Zheng Y, Li XB. *Arabidopsis* droughtinduced protein Di19-3 participates in plant response to drought and high salinity stresses. Plant Molecular Biology. 2014;**86**(6):609-625

*Plant Stress Physiology*

Science. 2018;**9**

gene, ZjZFN1, improved salt tolerance in *Arabidopsis*. Frontiers in Plant

*Arabidopsis* and rice. Plant Molecular Biology Reporter. 2010;**29**(1):185-196

Functional analysis ofTaDi19A, a saltresponsive gene in wheat. Plant, Cell &

[24] Jin Y-M, Piao R, Yan Y-F, Chen M, Wang L, He H, et al. Overexpression of a new zinc finger protein transcription factors CTZFP8 improves cold tolerance

[25] Rizhsky L, Davletova S, Liang H, Mittler R. The zinc finger protein Zat12 is required for cytosolic ascorbate peroxidase 1 expression during oxidative stress in *Arabidopsis*. Journal of Biological Chemistry. 2004;**279**(12):11736-11743

[26] Davletova S, Schlauch K, Coutu J, Mittler R. The zinc-finger protein Zat12

plays a central role in reactive oxygen and abiotic stress signaling in *Arabidopsis*. Plant Physiology.

[27] Zhang H, Ni L, Liu Y, Wang Y, Zhang A, Tan M, et al. The C2H2-type zinc finger protein ZFP182 is involved in abscisic acid-induced antioxidant defense in rice. Journal of Integrative Plant Biology. 2012;**54**(7):500-510

[28] Zhang H, Liu Y, Wen F, Yao D, Wang L, Guo J, et al. A novel rice C2H2-

type zinc finger protein, ZFP36, is a key player involved in abscisic acid-induced antioxidant defence and oxidative stress tolerance in rice. Journal of Experimental Botany.

[29] Yin M, Wang Y, Zhang L, Li J, Quan W, Yang L, et al. The *Arabidopsis* Cys2/His2 zinc finger transcription factor ZAT18 is a positive regulator of plant tolerance to drought stress. Journal of Experimental Botany.

2014;**65**(20):5795-5809

2017;**68**(11):2991-3005

2005;**139**(2):847-856

in rice. International Journal of Genomics. 2018;**2018**:1-13

[23] Li S, Xu C, Yang Y, Xia G.

Environment. 2010

[16] Ma X, Liang W, Gu P, Huang Z. Salt

tolerance function of the novel C2H2-type zinc finger protein TaZNF in wheat. Plant Physiology and Biochemistry. 2016;**106**:129-140

[17] Luo X, Bai X, Zhu D, Li Y, Ji W, Cai H, et al. GsZFP1, a new Cys2/ His2-type zinc-finger protein, is a positive regulator of plant tolerance to cold and drought stress. Planta.

[18] Huang J, Wang JF, Wang QH, Zhang HS. Identification of a rice zinc finger protein whose expression is transiently induced by drought, cold but not by salinity and abscisic acid. DNA

Sequence. 2009;**16**(2):130-136

[19] Chen F, Yu G-H, Jiang L-L, Ma X-F, Xu Z-S, Liu M-M, et al. A soybean C2H2-type zinc finger gene

GmZF1 enhanced cold tolerance in transgenic *Arabidopsis*. PLOS One.

[20] He F, Li HG, Wang JJ, Su Y, Wang HL, Feng CH, et al. Pe STZ 1, a C2H2-type zinc finger transcription factor from *Populus euphratica*, enhances freezing tolerance through modulation of ROS scavenging by directly regulating Pe APX 2. Plant Biotechnology Journal.

[21] Kim JC, Lee SH, Cheong YH, Yoo C-M, Lee SI, Chun HJ, et al. A novel cold-inducible zinc finger protein from soybean, SCOF-1, enhances cold tolerance in transgenic plants. The Plant

[22] Zhang X, Guo X, Lei C, Cheng Z, Lin Q, Wang J, et al. Overexpression of SlCZFP1, a novel TFIIIA-type zinc finger protein from tomato, confers enhanced cold tolerance in transgenic

2014;**9**(10):e109399

2019;**17**(11):2169-2183

Journal. 2001;**25**(3):247-259

2011;**235**(6):1141-1155

**198**

[31] Sakamoto H, Maruyama K, Sakuma Y, Meshi T, Iwabuchi M, Shinozaki K, et al. *Arabidopsis* Cys2/His2-type zinc-finger proteins function as transcription repressors under drought, cold, and high-salinity stress conditions. Plant Physiology. 2004;**136**(1):2734-2746

[32] Zhang X, Zhang B, Li MJ, Yin XM, Huang LF, Cui YC, et al. OsMSR15 encoding a rice C2H2-type zinc finger protein confers enhanced drought tolerance in transgenic *Arabidopsis*. Journal of Plant Biology. 2016;**59**(3):271-281

[33] Chang H, Chen D, Kam J, Richardson T, Drenth J, Guo X, et al. Abiotic stress upregulated TaZFP34 represses the expression of type-B response regulator and SHY2 genes and enhances root to shoot ratio in wheat. Plant Science. 2016;**252**:88-102

[34] Song B, Zhang Y, Li Y, Fu Y, Wang P. Novel drought-inducible Cys2/His2-type zinc finger protein STF-2 from soybean (*Glycine max*) enhances drought tolerance in transgenic plants. Pakistan Journal of Botany. 2019;**51**(3)

[35] Tran V-T, Braus-Stromeyer SA, Kusch H, Reusche M, Kaever A, Kühn A, et al. Verticillium transcription activator of adhesion Vta2 suppresses microsclerotia formation and is required for systemic infection of plant roots. New Phytologist. 2014;**202**(2):565-581

[36] Lawrence SD, Novak NG. Overexpression of StZFP2 in *Solanum tuberosum* L. var. Kennebec (potato) inhibits growth of Tobacco Hornworm larvae (THW, *Manduca sexta* L.). Plant Signaling & Behavior. 2018;**13**(7):e1489668

[37] Wang J, Hou Q, Li P, Yang L, Sun X, Benedito VA, et al. Diverse functions of multidrug and toxin extrusion (MATE) transporters in citric acid efflux and metal homeostasis in *Medicago truncatula*. The Plant Journal. 2017;**90**(1):79-95

[38] Tsutsui T, Yamaji N, Feng MJ. Identification of a cis-acting element of ART1, a C2H2-type zinc-finger transcription factor for aluminum tolerance in rice. Plant Physiology. 2011;**156**(2):925-931

**201**

**Chapter 11**

**Abstract**

using gene-specific markers.

breeding, phytohormones

**1. Introduction**

Genes for Different Abiotic

Stresses Tolerance in Wheat

*Sudhir Kumar, Shampa Purkyastha, Chandan Roy,* 

In the recent past years, global warming and climate change have drastically affected the agricultural crop productivity grown in tropical and subtropical areas globally by appearing to several new biotic and abiotic stresses. Among the abiotic stresses, heat, drought, moisture, and salt stresses are most prevalent. Wheat is the most common and widely used crops due to its economic and social values. Many parts of the world depend on this crop for food and feed, and its productivity is highly vulnerable to these abiotic stresses. Improving tolerance to these abiotic stresses is a very challenging assignment for wheat researchers, and more research is needed to better understand these stresses. The progress made in understanding these abiotic stress tolerances is due to advances in three main research areas: physiology, genetic, and breeding research. The physiology research focused on the alternative physiological and biochemical metabolic pathways that plants use when exposed to abiotic stresses. Identifying genes contributing to particular stress tolerance is very important. New wheat genotypes having a high degree of abiotic stress tolerance are produced through marker-assisted breeding by making crosses from promising concerned stress-tolerant genotypes and selecting among their progeny

**Keywords:** climate change, abiotic stress, wheat, physiology, genetic, marker assisted

Wheat is the second most important cereal crops of the world occupying about

220 million hectares area (mha) with a production of 716 million tons of food grain with a productivity of 3.2 tons per hectare [1]. It is extensively grown in Asia particularly in China and India. In India its production is enhances after the green revolution of late 1960s followed by another green evolution during 1980s. During these two green revolutions, the rate of annual growth in wheat production globally was ~3%, but in recent years it is declined to <0.9% due to appearance of new biotic and abiotic stresses. Although currently, the global wheat production has been able to meet the current demand and consumption, but we will have to enhance production and achieve the targets of at least ~858 Mt to meet the demand in 2050, as against current global production of 763 Mt. It comprises amounts to at least ~15% desired increase in global wheat production (1.5% annual increase)

*Tushar Ranjan and Rakesh Deo Ranjan*

### **Chapter 11**
