Role and Function of Eethylene Response Factor in Different Plants under Multiple Biotic and Abiotic Stresses

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Bibliographic Information
Journal Pakistan Journal of Biochemistry and Biotechnology
Title Role and Function of Eethylene Response Factor in Different Plants under Multiple Biotic and Abiotic Stresses
Author(s) Akram, Rabia, Farah Deeba, Maryam Zain, Nadia Iqbal
Volume 1
Issue 1
Year 2020
Pages 1-19
Full Text Crystal Clear mimetype pdf.png
URL Link
Chicago 16th Akram, Rabia, Farah Deeba, Maryam Zain, Nadia Iqbal. "Role and Function of Eethylene Response Factor in Different Plants under Multiple Biotic and Abiotic Stresses." Pakistan Journal of Biochemistry and Biotechnology 1, no. 1 (2020).
APA 6th Akram, R., Deeba, F., Zain, M., Iqbal, N. (2020). Role and Function of Eethylene Response Factor in Different Plants under Multiple Biotic and Abiotic Stresses. Pakistan Journal of Biochemistry and Biotechnology, 1(1).
MHRA Akram, Rabia, Farah Deeba, Maryam Zain, Nadia Iqbal. 2020. 'Role and Function of Eethylene Response Factor in Different Plants under Multiple Biotic and Abiotic Stresses', Pakistan Journal of Biochemistry and Biotechnology, 1.
MLA Akram, Rabia, Farah Deeba, Maryam Zain, Nadia Iqbal. "Role and Function of Eethylene Response Factor in Different Plants under Multiple Biotic and Abiotic Stresses." Pakistan Journal of Biochemistry and Biotechnology 1.1 (2020). Print.
Harvard AKRAM, R., DEEBA, F., ZAIN, M., IQBAL, N. 2020. Role and Function of Eethylene Response Factor in Different Plants under Multiple Biotic and Abiotic Stresses. Pakistan Journal of Biochemistry and Biotechnology, 1.

Abstract

These stresses effect crop production and quality, thus result is in economic lose and food insecurity. Many factors play vital role in regulating growth of plants along with developmental pathways during biotic and abiotic stresses. Transcription factors are proteins that control physiological, developmental and stress responses in plants. Ethylene response factors belong to the biggest family of transcription factors, known to participate in various stress tolerance like drought, heat, salt and cold. They are significant regulators of plant gene expression. The objective of this review is to present how ethylene response factor family proteins became the focus of stress tolerance as well as the development and growth of plants.

REVIEW ARTICLE

Role and function of Ethylene Response Factor in different plants under multiple biotic and abiotic Stresses

Rabia Akram1, Farah Deeba1, Maryam zain1, Nadia Iqbal1*

1Department of Biochemistry and Biotechnology, The Women University, Multan, Pakistan

*Corresponding Author:

Dr. Nadia Iqbal

Department of Biochemistry and Biotechnology, The Women University, Multan, Pakistan

Naaaadia7@gmail.com

Abstract

Abiotic and biotic stresses are the causes of drastic changes in plants growth and development.

These stresses effect crop production and quality, thus result is in economic lose and food insecurity. Many factors play vital role in regulating growth of plants along with developmental pathways during biotic and abiotic stresses. Transcription factors are proteins that control physiological, developmental and stress responses in plants. Ethylene response factors belong to the biggest family of transcription factors, known to participate in various stress tolerance like drought, heat, salt and cold. They are significant regulators of plant gene expression. The objective of this review is to present how ethylene response factor family proteins became the focus of stress tolerance as well as the development and growth of plants.

Introduction of ethylene response factor gene

Plants are susceptible to several types of environmental stresses throughout their life cycle. Theseadverse environmental conditions, specifically high salt concentration, temperature, soil moisture affect the growth, development, yield and geological distribution of cultivated crops across the world and decrease possible crop productivity approximately to 70% (1).

Biotic stresses for example weeds, diseases and insects occur at variable extent which may not be relevant in a particular developmental stage but decrease plant productivity (2).One of the main objective of cop breeding program is development of stress tolerant crop (3). Several widespread abiotic stresses reported to take place around the world are salinity (4), drought (5), water logging, extreme temperatures (6), also nutrient deficiency (7).

A variety of defense mechanism like maintenance of membrane integrity, generation of antioxidants, hormone regulation, stimulation of stress proteins and carbon fixation rate have been found to work in plant. Various mechanisms in plant are governed involving particular genes for these stresses(8) and help plants to protect against a series of outside stresses during morphological adaptation (9). A large number of genes with diverse functions are induced or repressed when a plant is undergo to abiotic stresses. These genes encode for proteins which could be divided into two groups. One of them is primary group comprises functional proteins that is key enzymes for osmolytes biosynthesis like proline and sugar alcohols, molecular chaperones, and membrane transporters which are directly related to defense of plants from abiotic stress. The other groupcontains proteins that are regulatory in nature and control signal transduction and stress receptive gene expression and consists of a variety of transcription factors and protein kinases (10).

Transcription factors (TFs) that are regulatory proteins have a central part in initiatingdefense gene expression (11). They interact throughcis-acting components present in promoter region of various stress regulated genes and consequently start cascades or entire system of genes that perform together increased tolerance to various stresses at a time. Stress responsive transcription factors are strongtools for genetic engineering because their overexpression can result in either upregulation or downregulation of entire collection of genes under their control.A large number of transcription factors participate in plant stress tolerance and regulate plant responses to diverse stresses.InArabidopsis more than 1500 genes have reported to encode a various transcription factors(12). Some of the transcription factorsrelated to plant stress response are categorized into numerous large families for example AP2/ERF, NAC, bZIP, zinc-finger,MYB, MYC and WRKY (13).Eethylene responsive factoris the member of large gene family the APETALA2/Ethylene-Responsive Factor superfamily. This superfamily also include AP2as well as RAV family(12). This superfamily has characteristic AP2/ERF domain, which contains near about sixty to seventy amino acids. The families of AP2/ERF superfamily defined like, proteins of AP2 family have AP2/ERF domains which are two repeated domains, and proteins of ERF family consist of one domain that is AP2/ERF, whereasproteins of RAV family have a B3 domain. The ethylene response factor family isfurther divided into two distinctive subfamilies, one is CBF/DREB subfamily, second is ERF subfamily (14). The ERF region initially identified as conserved element in four proteins characterized in Nicotiana tabacum. These proteins are distinctive ethylene responsive element binding proteins, named ERF 1, 2, 3, 4 also found to interact with consensus promoter sequence GCC box which play role in ethylene responsive transcription of genes (15). The ERF subfamily proteins were classified into six sets, B-1 to B-6. Nakano et al., (16) studied the expression of various genes of ERF subfamily in various plants and their biological roles.

The ethylene responsive factors also governed by feedback mechanism and may be ethylene dependent or ethylene independent. Apart from regulation mechanism, ERFs expressions are regulated post-transcriptionally via micro RNAs, while expressions of miRNA are also regulated through ERFs. Ethylene responsive factors have found to involve as activator or various genes while it may act as repressor of various genes during of plants defense response averse to biotic stress condition (17). Ethylene responsive factors also defense role in fungal and bacterial diseases responses as well as during viral infection in plants (18). SERF1 overexpression enhanced tolerance of salinity through ROS stimulated MAPK indicating flow (19). TaERF1 in wheat was appeared in the direction of improve easiness to abiotic, also biotic stress conditions (20). AP2/EREBP family which are plant-specific TFs exposed by Arabidopsis mutational analysis, that were liable for enhance wax substance, decline stomatal density, changed cuticle premises in addition to the connected important raise in tolerance of drought Arabidopsis modified shine (21).

Ethylene response factors (ERF) are associated in stress tolerance in a number of species of plants(22), it was revealed that GhERF2, GhERF3 and GhERF6 can proceed as significant frequent components of various signaling pathways receptive to abiotic and biotic stresses. To advance identify with their activity, the development of transgenic plants overexpressing these genes is ongoing (23).

Discovery

ERFs previously named ethylene response binding proteins denoted by ERE binding proteins. The ethylene responsive element proteins earliest extracted from plant Nicotiana tabacum(15). ERFs consist of a domain, AP2 DNA-binding region, as of a plant-definite fabulous family of 122 TFs in Arabidopsis. In genome of tomato, near 85 gene of ethylene responsive factors family protein stay uncharacterized (24). It is described that in various plants ERF domains product be present within several regulatory genes (25, 26) but is not present in fungi, yeast or mammalian (27). In the most recent decade, to a large extent, consideration have been accentuated on AP2/ERF large family. The hypothesis elucidates as the lateral gene movement of HNH-AP2 endonuclease originated from bacteria or virus into the host plant might bring about regenerate the AP2/ERF great family (28).

Functional diversity in plants

Up to now, Ethylene responsive TFs have been recognized as well as distinguished from numerous species of plants for example Arabidopsis (16), poplar (29), cotton (30), soybean (31), barley (32), maize (33), cucumber (34), apple, wheat (35), rice (36),sorghum (37), Medicago truncatula (38), also potato (39). Overall 147 genes in Arabidopsis, encoding AP2/ERF TFs, transcription factors named ERF encoded by 122 of these 147. ERF TFs of Arabidopsis may be divided into 12 groups different from each other, found on phylogenetic analysis, these groups named as I to X, VI-L,Xb-L (16).Functional diversity of ethylene response factor in different plants is given in table 1.

Structure

The ERF TFs take part in plants development procedures, in response to environmental stresses and hormone signaling (39). The ethylene response factor family has been comprehensively studied in recent times. In AP2/ERF domain b-sheet, 2 conserved amino acid residues distinguished ERF transcription factors, residues are 14th Ala as well as 19thAsp (14).ERF TFs allocate a conserved 58–59 amino acid domain (the ethylene response factor domain) and encompass sturdy capability to bind an extensive s rt of CREs within promoter of mark genes. From the cis-regulatory elements, DRE/CRT and GCC-box are the two chief DNA-binding elements (40). Most stimulatingly, numerous studies expose ERF proteins interconnect by DRE/CRT motif, a cis-acting element which acts in response to osmotic stress or cold (41), they build up defense by inter relating among stress-regulated genes (42).

The α-helix as well as β-sheet of ERF region identify the GCC box (called core cis-acting element AGCCGCC) of objective DNA on the ER promoter region (43). Pathogenesis-related (PR) genes are expressed by these GCC box elements (44). Additionally, the minority conservative elements of other than AP2 region grant transcriptional repression or activation in favor of particular gene, for example ERF connected amphiphilic oppression element that is EAR:DLNxxP (45), LWSY element (30)in addition to B3 oppression domain motif that is BRD:RLFGV (46). ERF protein either has homology resemblance with identified DNA joining proteins nor has fundamental zinc finger or leucine zipper elements, which propose ERF is a set of novel DNA binding proteins (43).Figure 1 gives classification and highlights functional association among AP2/ERFs in a variety of abiotic stress responses plants.

Role of ethylene response factor

Ethylene responsive factors take part essential regulatory tasks in various developmental processes and stress responses in plants. Genes of ERF family in an extensive array of plant types, were categorized into 8groups and 14 subgroups on the base of gene structures, conserved motifs, phylogenetic relationships, and biological functions. ERF genes, particularly in groups II as well as VII, may be utilized like contenders to develop crop resistance, for the reason that overexpression of ERF genes increases various disease resistance along with progresseasiness to drought, salt, freezing also in transgenic plant. The inclusive testing of phylogenetic correlations, conserved pattern, moreover physiological functions is beneficial in illuminating the biological roles of genes of ERF family in signal transduction, gene regulation along with defense response below stress environments (11).

ERF TFs have been revealed to be concerned in the regulation of expression of lipids, cell wall constituents, pathogenesis related genes, biosynthesis genes, basic type defense related genes, chitinase, osmotin, moreover b-1,3-glucanase encoding genes (28). Few of them are exposed to be concerned with in the control of plant reactions to biotic, also abiotic pressures by either activating or repressing abscisic acid responsive genes (47). Such as AtERF4 over-expression plant was a smaller amount of responsive to ABA suppressed root enlargement that entails negative control of ABA also ethylene reactions (48). The expression of ABA-responsive genes are repressed by AtERF7 that binds to the GCC box (49).ERF111 or ABR1 takes part as a negative controller of ABA reactions throughout seed germination, ABA and stress regulated gene expression (50) wherever as in Arabidopsis ABA hypersensitivity is conferred by transgenic plant, overexpressing AtERF13 (51). AtERF15 exposed on the way to proceed as a activist controller of ABA reactions (52). In contrast, ABA may also bring about the few ERF genes expression such as the expression of tomato ERF gene JERF1/3, cotton ERF gene GbERF in addition to tobacco ERF gene NtCEF1 has been revealed to be evoked by ABA (53).Sub group IX in IXERF subfamily accommodates 4 tiny ERFs through amino acids, varied as of 131-139. These tiny ethylene response fators named ERF95, ERF96, ERF97,ERF98. Additionally toward AP2/ERF area, these ethylene response factor have unfamiliarrole pattern termedCMIX-1 (16). In the group of these, ERF95 in addition called ethylene and salt inducible1 (ESE1), furthermore ERF98 have been exposed to concerned with the control of salt acceptance (54). In the past ERF97termed AtERF14, it has been revealed to control plants protection retort (55). In recent times ERF96 also has been revealed to regulate plant defense response (56).

Role of ERF during Stress

ERFs are concerned during the parameter of plant growth, expansion, metabolism as well as plant reactions to ecological stimulus like abiotic and biotic stresses (57). The ERF transcription factors are participated in the ethylene signaling pathway as well as directing in the direction of the expression of a variety of protection-associated genes, for example pathogenesis-related genes (40), abiotic stress receptive genes (58). The existence of a GCC-box or a dehydration responsive element/C-repeat element (DRE/CRT, CCGAC) situated in the promoter sequences of defense-related genes are involved in characterization of these genes (14). Figure 2 shows stress related genes expression, their role under stresses

Table 1: Examples of ERF family proteins involved in regulation of multiple abiotic stress responses in different plants.


Figure 1: AP2/ERF transcription factors in plants, their classification and their functions has also been reported.

Role of ERF during Stress

ERFs are concerned during the parameter of plant growth, expansion, metabolism as well as plant reactions to ecological stimulus likeabiotic and biotic stresses (57). The ERF transcription factors are participated in the ethylene signaling pathway as well as directing in the direction of the expression of a variety of protection-associated genes, for example pathogenesis-related genes (40), abiotic stress receptive genes (58). The existence of a GCC-box or a dehydration responsive element/C-repeat element (DRE/CRT, CCGAC) situated in the promoter sequences ofdefense-related genes are involved in characterization of these genes (14). Figure 2 shows stress related genes expression, their role under stresses.

Abiotic stress

Environmental stresses that are nutrient deficiency, drought, salinity, extreme temperatures and chemical toxicity, leads to a chain of physiological,morphological, molecular, and biochemical changes which negatively have an effect on plant development, yield and survival all-inclusive (59); (60), in responseto those stressesplants develop mechanisms. Plant feedbacks are multifarious, including extensivemetabolic and transcriptional events (61). The molecular and physiological studies of environmental stresses have proposed a familiar network of several signaling tracks that mediate salt, extreme temperatures and drought stress responses in plants (62). In abiotic stress responses the DREBs proteins are most extremelyconsidered ethylene response factor. Elements of the subfamily DREB1/CBFs are quickly induced in retort to chilly stress andget better tolerance to freezing when expressed ectopically, (63). Whereas,RNAi lines of Arabidopsis for CBF1 or CBF3 have decreased cold tolerance (64). The very fact, for freezing tolerance CBF2associated by quantitative trait locus (65), point out that DREB1/CBF genes are cold activated and in Arabidopsis are liable to be mainmonitor of reaction to wintry stress. Furthermorein Arabidopsis CBF2 abnormal gene communication has been revealed to retard leaf senescence, indicating that CBFs prop upchillysurvivalthrough deceleration down growth moreover delaying blossoming in coldnesstill temperatures raise in spring (66).

Figure 2: A representive model of process of abiotic or biotic stress tolerance in plants.

The subgroup of DREB2 contains eight elements in Arabidopsis, moreoverhomologs in the genetic material of numerous angiosperm families. Along with the components of the DREB2 subdivison, Arabidopsis DREB2A as well as DREB2B were induced in loss of water, heat and elevated saltiness in an ABA independent way (67). The abnormal gene expression of a constitutively dynamicstructure of DREB2A demonstrates developed forbearance to famine, heat and elevated salt stresses (67);(68) whereas DREB2A are furthersusceptible to temperature Shock (67). From rice or Arabidopsis several ERF-VII genes areexposed to be participated with in response to hypoxia as well assubmergence (69).In Arabidopsis RAP2.12 are constitutive ERF-VII factors, they arerecommended to proceed as prime triggers for the molecular feedback to insufficiency of oxygen (70); (71), that is then comforted at the oxygen deprivation activated ethylene response factors HRE1 also HRE2 (72). In submerged deepwater rice, SK1and SK2 motivate internodes elongation to effect clearance of the water level (73), whereas Sub1A promotes a quiescent strategy that allows carbohydrate saving and improves tolerance after flash-flood (74). A number of ERF genes from various plants have been exposed to confer various stress tolerance when expressed ectopically (75). This ‘unspecific’ effect may be explained by the activation of tolerance pathways whichreduce a generic stress statusfor example oxidative bursts generated as results of the primary stresses. Otherwise constitutive ERF expression could set the plant in a common alert state that fasten or expand the response when a specific stress is applied (28).

Biotic stress

ERFs are entailed inbiotic stress responses, ethylene response factor proteins initiallyextracted the same as transcription factors (TFs) whichconnect to promoter regions of stress-responsive genes. ERF genes assessed to date, are provoked through biotic stresses, consist of wounding, germ contagionstresses, such as transcription of basic type pathogenesis-related (PR) genes, defense-related genes,chitinase, osmotin, and b-1,3-glucanase activated by numerous ERFs. On the other hand, the group of target genes regulated by each ERF has not been entirelyrevealed. ERF1 as well as its homologs be associated to ERF-IX group, are possibly the ERF transcription factors whose participation in pathogen reaction in Arabidopsis has been most widely distinguished (76), as wella number ofproteins of ERF adjust ethylene biogenesis (77). Tobacco OPBP1 when expressed ectopically in transgenic rice increases resistance to pathogens (78).

Figure 3: Hormonal signalling and expression of genes in biotic stresses.

Conclusion

The existing information about various roles ofAP2/ERFs recommends that the members of this large superfamilycan be utilized for improving stress tolerancefor better crop yields. The members of this family can be employed in improving nutritional quality of grown varieties. Up to now extensiveadvancement has been done to know the functions ofAP2/ERFs,in response to various stresses. To develop multiple stress tolerant crops the members of this superfamily have been used.The plants genome sequencing projects have facilitated for recognition of ethylene response factors in different plants. It is stated that the proteins of ERF family has ability to confer resistance against numerous stresses in crop plant.

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References