Extensive intragenic divergences amongst ancient WRKY transcription factor gene family is largely associated with their functional diversity in plants

Journal Pre-proof Extensive intragenic divergences amongst ancient WRKY transcription factor gene family is largely associated with their functional diversity in plants

Aniruddhabhai Khuman, Sakshi Arora, Harshita Makkar, Abhishek Patel, Bhupendra Chaudhary PII:

S2352-4073(20)30003-2

DOI:

https://doi.org/10.1016/j.plgene.2020.100222

Reference:

PLGENE 100222

To appear in:

Plant Gene

Received date:

26 September 2019

Revised date:

20 December 2019

Accepted date:

24 January 2020

Please cite this article as: A. Khuman, S. Arora, H. Makkar, et al., Extensive intragenic divergences amongst ancient WRKY transcription factor gene family is largely associated with their functional diversity in plants, Plant Gene (2019), https://doi.org/10.1016/ j.plgene.2020.100222

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© 2019 Published by Elsevier.

Journal Pre-proof Extensive intragenic divergences amongst ancient WRKY transcription factor gene family is largely associated with their functional diversity in plants

Aniruddhabhai Khuman, Sakshi Arora, Harshita Makkar, Abhishek Patel and Bhupendra Chaudhary* School of Biotechnology, Gautam Buddha University, Greater Noida, India-201310 *Corresponding author: [email protected]

Abstract

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Diversity at gene sequence and function levels is an evolutionary outcome of a constantly

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expanding gene family in plants. Genome sequences have provided a platform to comprehend the evolutionary architecture of genes and gene families, especially in plant genera.

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Apparently, regulatory sequences and the selection of trans-binding proteins have massive impressions of gene expression evolution as compared with a transformation at genic-level

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itself. Therefore, it becomes essential to comprehensively dissect the genome-wide structure-

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function evolution of regulatory regions’-binding trans-proteins. WRKY gene family members are ancient and developmentally and functionally diverged transcription factors that notably regulates numerous cellular as well as stress responsive phenotypes in plant species.

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To comprehend the evolutionary consequences of WRKY genes during plant development, a meta-phylogenetic analyses of at least 266 WRKY amino acid sequences of alga, monocot

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and eudicots plant species was accomplished. Phylogenetic clustering identified assorted groups and sub-groups of WRKY orthologous or paralogous sequences. Paralogous gene divergences of monocot and eudicot species showed strong ‘purifying’ selection during the sequence

evolution.

Evidently,

evolutionary

exon-intron

structural

diversification

predominantly arose by ‘exon/intron-gain’ and insertion/deletion during primeval sequenceexonization. The structural DNA-binding domain, and global structures of WRKY proteins are exceedingly divergent and also explicit to the classified phylogenetic clusters. Expression profiling displayed that GroupIIe WRKY members are profusely expressed in root tissues of the representative species. Remarkably, the spatial and temporal gene expression patterns of WRKY genes in plants apprise our perception about their contributions in the regulation of cellular functions; and structural-functionalism of these members speculate their probable employment into future crop improvement programs. Keywords: WRKY, phylogeny, expression profiling, exon/intron, plants

Journal Pre-proof Introduction Plants are continuously exposed under biotic or abiotic stress that impact directly on the productivity of economically important crops. In order to cope up with environmentally induced stress conditions, plants have developed various mechanisms regulated by various genes and regulatory factors are differentially expressed (Dou et al., 2014). For differential expression of these genes various signaling cascades are employed consisting of transcription factors, regulatory molecules like microRNAs, tasiRNAs, and signaling molecules including various types of kinases (Phukan et al., 2015). Transcription factors (TFs) are very crucial in the developmental or stress-induced regulation of signaling cascades or mechanisms. These

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factors regulate expression of various genes by binding to their promoter regions (Phukan et

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al., 2015). TFs also act as an activator or repressor of any particular gene(s) expression under different environmental conditions (Song and Gao, 2014). MYB, NAC, ERF, SRS, WRKY are

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selective examples of stress-induced transcription factor reported to be present in plant cell

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(MacMillan et al., 2017).

WRKY transcription factor gene family consists of several gene members amongst plant

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species with highly conserved WRKYGQK domain (Eulgem et al., 2000). The number of WRKY gene family members drastically vary among different plant species, for example, 100

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gene members in Oryza sativa, 75 members in Arabidopsis thaliana and 116 members in Gossypium hirsutum (Dou et al., 2014). WRKY gene family members are spatially and

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temporally regulated and play important roles in plant development, seed germination, biotic and abiotic stress responses (Scarpeci et al., 2013). In Arabidopsis, AtWRKY6 and AtWRKY42 have been reported to play important roles in response to low Pi-stress by regulating PHO1 gene expression (Chen et al., 2009). AtWRKY42 modulated Pi-homeostasis by regulating the expression of PHO1 gene (Su et al., 2015). AtWRKY45 is involved in Pi-starvation by direct up-regulation of phosphate transporter PHT1.1 gene expression (Wang et al., 2014a). AtWRKY12 and AtWRKY13 have similar sequences and are involved in the regulation of flowering time phenotypes under short day conditions (Li et al., 2016). AtWRKY18 and AtWRKY60 show positive effects on ABA sensitivity for inhibition of seed germination and root growth (Chen et al., 2010). AtWRKY22 is involved in the regulation of leaf senescence under dark conditions (Zhou et al., 2011). The over-expression of AtWRKY25, AtWRKY26 and AtWRKY33 led to an increase in the heat shock protein (hsp) related signaling pathway and enhanced resistance to heat stress in Arabidopsis (Li et al., 2011). AtWRKY34 and AtWRKY2 play an important role in MPK3/MPK6 cascade during pollen development in

Journal Pre-proof Arabidopsis (Guan et al., 2014). Increased expression of AtWRKY53 regulates stomatal movement via reduction of H2O2 content and promotion of starch metabolism in the guard cells thus having crucial role during drought tolerance (Sun and Yu, 2015). AtWRKY54 and AtWRKY70 negatively regulate stomatal closure and consequently is involved in osmotic stress tolerance (Li et al., 2013). In rice, OsWRKY53 and OsWRKY71 are involved in elicitorinduced defense signaling pathway (Chujo et al., 2007; Chujo et al., 2008). OsWRKY17, OsWRKY23 and OsWRKY82 genes are involved in induced leaf senescence in rice (Liu et al., 2008; Sperotto et al., 2008; Jing et al., 2009). Similarly, in cotton, GhWRKY25 is reported to regulate various abiotic stress responses and other multiple defense mechanisms (Liu et al.,

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2015). Overexpression of GhWRKY40 down-regulates defense-related genes and enhanced

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wounding tolerance and susceptibility towards the Ralstonia solanacearum (Wang et al., 2014a). Also, GhWRKY15 is involved in disease resistant and plant development (Yu et al.,

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2012). On contrary, GhWRKY33 act as negative regulator to drought stress and participate in ABA signaling pathway (Wang et al., 2019). Remarkably, WRKY transcription factors play

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essential roles in plant growth and development as well as in stress-related phenotypes

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mainly through the tight regulation of cell-signaling networks. Currently, the plant genome sequencing databases provide incredible evidences about the

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genes, regulatory functions; and their evolutionary imprints corresponding to the evolution of plant kingdom. Diversification of gene sequences at evolutionary scale is crucial and set

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examples for the functional expansion of gene members preferentially during selection of superior phenotypic traits amongst crop plant species. Here, we aim to identify WRKY gene sequences from representative monocot and dicots plant species and establish their evolutionary association, sequence variability, and evolution of WRKY motif with their functional attributes at system level. Spatial- temporal expression represent that patterns of structural divergences with functional conservations of WRKY genes are largely speciesspecific and not global. Presented results are novel and underscore the evolutionary trajectories of WRKY genes in plant kingdom and recommends their field employment for future crop advancement. Materials and methods Identification of WRKY gene family members amongst plant species Different species specific WRKY gene sequences were obtained from multiple database including National Centre for Biotechnology Information (NCBI); plant transcription factor

Journal Pre-proof database (http://planttfdb.cbi.pku.edu.cn/); Arabidopsis information resources (TAIR) database

(https://www.arabidopsis.org/);

rice

genome

annotation

project

(RGAP)

(http://rice.plantbiology.msu.edu/); cotton genome database (https://www.cottongen.org/) (S1 Table). Amino acid sequences of WRKY gene family members from lower plant (green algae), monocot (rice), and dicots (Arabidopsis and cotton) plant species were downloaded from the NCBI (https://www.ncbi.nlm.nih.gov). The WRKY gene family amino acid sequences subsequently used to identify various WRKY homologues and paralogous sequences of sequenced-genome resources through BLASTP. The e-value for BLASTP was

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set at 1e-10 to obtain the final dataset of a WRKY protein.

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Multiple sequence alignment and phylogenetic analysis

Multiple sequence alignment of all conserved WRKY domains was determined by using

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Clustal Omega software (https://www.ebi.ac.uk/Tools/msa/clustalo/) with default parameters. The Bayesian Algorithm of phylogenetics was employed and protein sequences were

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submitted into Interactive Tree of Life (iTOL) web server for the construction of

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phylogenetic tree.

Estimation of WRKY protein conservation and Ka/Ks analysis

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WRKY protein conservation at evolutionary scale was estimated using ConSurf Server (http://consurf.tau.ac.il/). To identify structural conservation of WRKY proteins, multiple

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sequence alignment files were submitted to ConSurf server. Calculations of conservation scores for each specific position of amino acid residues by means of an empirical Bayesian algorithm. Amino acid residues were clustered at average, variable and conserved scale for three-dimensional visualization purpose. Non-synonymous (Ka) and synonymous (Ks) substitution

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each

gene

pair

was

calculated

using

PAL2NAL

(http://www.bork.embl.de/pal2nal/) program. In this program, following commands were performed- Codon table  universal; Remove mismatches  yes; Use only selected positions 

yes; Output format  PAML. With these settings, the codon alignment corresponding to

the specified regions is generated in the PAML format. Structural organization and divergences of exon-intron regions The exon-intron organization pattern of WRKY genes was generated using online Gene Structure Display Server (GSDS 2.0) (http://gsds.cbi.pku.edu.cn/). To study the evolutionary trends of exon-intron organization of WRKY genes in plantae kingdom, the gene structural

Journal Pre-proof architectures were compared using Plant Intron Exon Comparison and Evolution database (PIECE)

online

tool

and

visualized

with

EXALIGN

viewer

(https://wheat.pw.usda.gov/piece/). Gene sequences were considered paralogous if e-value of Multiple Sequence Alignment (MSA)-based %sequence homologies among all gene members is equal or less than 1e-20. Evolution of “binding motif” of WRKY genes The conservation of amino acid sequence logo constructed using WebLogo3 online tool (http://weblogo.threeplusone.com/). Each of the amino acid sequences logo consists stacks

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letters, the letter height denotes the conservation of amino acid at any particular position. The sequence conservation at any position was measured in bits. Structural homology-modeling

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were performed using Swiss-model server (https://swissmodel.expasy.org/). Evolutionary

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conservation of WRKY domain and structural comparisons of Arabidopsis, rice, cotton and

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green algae were performed.

Expression analysis of WRKY genes in plants

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Expression data of different tissues of Arabidopsis, rice and cotton plants were retrieved from Transcription Variation Analysis (TRAVA) (http://travadb.org/); Rice Expression Database (http://expression.ic4r.org/);

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(RED)

and

GEO

dataset

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NCBI

(https://www.ncbi.nlm.nih.gov/pubmed) databases (S2 Table). To further analyse and

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visualization of large multi-dimensional dataset, the normalized expression data of various WRKY gene family members was utilized to generate the heatmap using Multi experiment viewer (MeV) software.

Results Evolution classification of WRKY proteins identifies conserved groups amongst monocot and dicot plants To study the evolutionary origin of WRKY gene family in plants, we analyzed WRKY gene family homologue and paralogue sequences across the species of green algae and plantae, which are believed to share conjoint ancestral node. At least 266 WRKY amino acid sequences, comprising of 86 sequences from monocot (Oryza sativa), 179 sequences from

Journal Pre-proof dicots (Arabidopsis thaliana and Gossypium hirsutum), and 1 lower-plant (green algae) were retrieved on the basis of their evolutionary association. These 266 WRKY sequences were aligned using Clustal Omega and the alignment results were subsequently used to construct a phylogenetic tree. Surprisingly we observed WRKY gene family members were evolutionary conserved (Song and Gao, 2014), and could be categorized into 3 major groups and sub-groups (Fig. 1). Interestingly, the phylogenetic result accentuated high diversity in members belonging to Group I and Group IIc, consisting members from the lower plants, monocots and dicots. The data indicates that WRKY protein

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sequences are ancient and conserved amongst the species and have evolved prior to any lineage diversification in nature. Group II members appeared into three distinct clades that is

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Group IId + IIe, IIc and Group IIa +IIb. Interestingly, Group IId and Group IIe shared a common ancestral origin (Fig. 1) (Zhang and Wang, 2005). The Group IIc is the largest

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group identified with 18 monocots (rice) and 48 dicots (cotton and Arabidopsis) members

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sharing a common ancestry.

Evidently, members of Group I contain two WRKY domains whereas members of other

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groups showed only one WRKY domain. It indicates that specific group/subgroup WRKY gene sequences are evolving independently and parallelly within a species and may be the result of

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an adaptive selection of such sequences at evolutionary tempo. Remarkably, WRKY gene of green alga was present in Group I of phylogenetic tree comprising of one WRKY domain.

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This indicated that Group I in modern plant species evolved from green algae, followed by ensuing homologous and/or paralogues genic-duplication or whole genome duplication (WGD) events followed by millennia of human selection, at least within domesticated plant species. From the evolutionary perspective, it emerges that the evolution of WRKY genes has occurred considerably prior to the divergence of ancient plant species into modern monocots and dicots species. GhWRKY18-OsWRKY53 (Group I), GhWRKY39-OsWRKY11 (Group IIc) and GhWRKY98-OsWRKY56 (Group IIc) largely predominates to be the noticeably conserved WRKY sequences (Fig. 1). These monocots and dicots species share a common node and considered as homologous sequences. Two dicots (AtWRKY55 & GhWRKY101) WRKY genes represent a clade with monocot sequence OsWRKY75 and deliberated as conserved orthologous sequences. These results showed analogous patterns of the evolution of WRKY gene sequences amongst dicot and monocot plant species, and suggest that WRKY paralogs are profusely diverged than their respective orthologs. Phylogenetic study suggested that the

Journal Pre-proof evolutionary classification of WRKY genes amongst monocot and dicot lineages is preserved; and commends the ancient evolution of WRKY gene family. Since the ancestral WRKY amino acid (aa) sequences of different kingdoms diverged from a common ancestor several million years ago, WRKY aa sequences of green algae, monocot (O. sativa) and dicot (A. thaliana and G. hirsutum) plant species were investigated to determine their evolutionary resemblances (Fig.2). This monocot and dicot plant species showed high homogeneity in WRKYGQK (Tryptophan, Arginin, Lysine, Tyrosine, Glycine, Glutamine, Lysine) domain as compared with algal domain except some variability at 6th

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amino acid position which is represented as glutamine in higher plants (Eulgem et al., 2000). Evidently, evolutionary divergence among WRKY sequences is prominent from lower plant

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to the higher plants, for example, green algae have solitary WRKY gene sequences whereas a

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large number of WRKY genes are present in monocot and dicot plant species.

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Evolutionary structural conservation of WRKY gene family in plants On basis of sequence homogeneity and phylogenetic analysis, 20 genes pairs were identified

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with 09 pairs of cotton and rice paralogs each; and 02 pairs of Arabidopsis (Table 1). The highest and the lowest amino acid sequence similarities identified amongst cotton were

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between GhWRKY14-GhWRKY17 (80.74%) and GhWRKY69-GhWRKY84 (71.81%), respectively. The highest and lowest amino acid sequence similarities identified amongst rice

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gene pairs were between OsWRKY61-OsWRKY81 (95.52%) and OsWRKY38-OsWRKY53 (74.62%), respectively. The highest and the lowest amino acid sequence similarities identified amongst Arabidopsis gene pairs were between AtWRKY64-AtWRKY67 (73.9%) and AtWRKY63-AtWRKY64 (73.02%), respectively. Using PAL2NAL program (Goldman et al., 1994), number of synonymous sites (S) and non-synonymous sites (N) present in the amino acid sequences were calculated. The synonymous (dS) and non-synonymous (dN) substitution rates were calculated using codon alignment results. It is evident that WRKY gene family amongst plant species has been evolutionary selected through the purifying selection nucleotide substitution rate ratio (dN/dS<1). The results suggest that WRKY paralogs gene pairs in monocot and dicot species have undergone strong purifying selection pressure during the sequence evolution (Table1). The patterns of evolutionary conservation of amino acid homologous and orthologous sequences were traced using site-specific scoring calculated by Bayesian algorithm of phylogenetics. In this study, specific regions were identified having high degree conservation in the WRKY domain of monocots and dicots species. Based on

Journal Pre-proof site specific scoring, seven amino acids- WRKYGQK long motif was identified in all representative species. Targeted amino acid positions of WRKY domain in these proteins showed maximum evolutionary conservation (Fig. 3). Remarkably, WRKY domain is conserved at the evolutionary scale ranging from lower plant species to higher monocots and dicots plants. Exon-intron organization of WRKY genes in plantae kingdom The exon-intron organization of WRKY gene family of dicot plant species were investigated by the comparison of genomic DNA sequences with their respective cDNA sequences using

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Gene Structure Display Server (GSDS). The exon-intron structure provides advanced knowledge of evolutionary classification of WRKY gene family in plants. It was observed that

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the number of exon and intron varied among WRKY gene family members and also between

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paralog member of both cotton and Arabidopsis (AtWRKY26 & GhWRKY18) (Fig. 4). Most of the AtWRKY and GhWRKY gene family members have 2-4 intronic regions present in their

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gene sequences. Surprisingly, AtWRKY19 gene has 15 intronic and 16 exonic regions whereas GhWRKY25 has 22 intronic and 23 exonic regions. Moreover, both these genes has the

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longest sequence amongst the respective gene family members. Among Arabidopsis WRKY gene family members, 52.85% genes have 2 intronic and 3 exonic regions, 15.71% genes

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have 3 intronic and 4 exonic regions and 11.42% genes have only 1 intronic region. GhWRKY116 has the longest intronic region between second and third exon and a total of 3

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exons and 2 introns. Large variations in the intronic and exonic length between ortholog and paralog across dicot species suggested that the evolution of exonic and intronic regions occurred after the monocot and dicot divergence. Structure of WRKY gene family of eudicots and monocot plant species were derived by using Plant Intron Exon Comparison and Evolution (PIECE) database. OsWRKY24 was used as representative WRKY and for studying exon and intron structural divergences occurred during WRKY gene evolution. The representative exonic sequence was aligned with other representative monocot and dicot species. Through this sequence alignment, precise matches and mismatches, as well as intron gain/loss events could also be determined. In result, the first exon shared the same frame but showed match-to-mismatch with aligned exonic region whereas second exon showed match-to-mismatch but did not shared a common frame. Also, second exon adhered to its length of 175-178 bp. The second exon of Arabidopsis showed exact match with Citrus clementine, and second exon of Medicago truncatula showed exact

Journal Pre-proof match with second exon of Glycine max (Fig. 5). These data indicate that exon-intron structure diversification occurred prior to the divergence of higher plants in plantae kingdom. Preservation of amino acid and ‘degree of nativeness’ amongst WRKY proteins WRKY transcription factors have diverse functions mainly in the regulation of plant growth and development at different developmental stages. To determine the evolutionary footprints of WRKY genes, sequence homology between 266 WRKY genes from G. hirsutum, O. sativa, A. thaliana and C. reinhardtii were investigated using multiple sequence alignment and WebLogo 3 online tool. In result, it was observed that 7 amino acid residue (WRKYGQK)

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was conserved between dicot species A. thaliana and G. hirsutum, monocot species O. sativa and green algae (C. reinhardtii) except 6th amino acid position which showed hyper-variable

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hydrophilic side chain containing glutamine or lysine residue. Out of these two amino acid

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residues, glutamine residue was more abundant in WRKY protein sequences in comparison

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with lysine residue (Fig. 6).

We predicted WRKY domain comparative 3D structures of representative species member of

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Arabidopsis, O. sativa, G. hirsutum and C. reinhardtii (Fig.7). These closely related representative sequences of WRKY domain structures are predicted using QMEAN

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(Qualitative Model Energy Analysis) method in SWISS-MODEL server. Dark blue color show high quality of model and orange color show least quality of model, this quality define

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through QMEAN Z-score. The QMEAN score is deriving through the ‘degree of nativeness’ of the predicted 3D structure (Benkert et al., 2010). These result showed evolutionary conservation of WRKY domain in the lower plant, monocots and dicots species. Expression profiling reveal structure-function relationship amongst WRKY genes The expression data of WRKY gene members in different tissues of three different species (A. thaliana, O. sativa and G. hirsutum) which represented evolutionary conserved Group IIb and Group IIe (Fig. 1) were obtained using TRAVA database, Rice expression database and NCBI, respectively. The WRKY expression data in different tissues such as leaves, anther, root, and seeds from dicot and monocot representative species were analyzed and visualized using heat map. It was observed that all WRKY members from the mentioned evolutionary conserved group were highly expressed in the root tissues (Fig. 8A). The alignment score of all WRKY genes that were expressing in roots of Arabidopsis, rice and cotton is less than 40, highlighting that the structurally diverged genes amongst various monocot and dicot plant

Journal Pre-proof species have conserved expression patterns; with preferably conserved functions. Whereas AtWRKY16 showed significant expression in all five developmental stages- root, leaves, anther, seed development and meristem. These results indicate that AtWRKY16 has constitutive expression patterns and may be attributing to the generalized functions at cellular level. On contrary, OsWRKY12, OsWRKY37, OsWRKY39, OsWRKY66, GhWRKY48 and GhWRKY58 were observed to be maximally expressed in roots and considered largely as root-specific genes in rice. GhWRKY57 and GhWRKY58 are evolutionary conserved and share a conserved node (Fig. 1). Similarly, OsWRKY31 and OsWRKY39 share common evolutionary conserved node but vary significantly in their spatial expression patterns (Fig. 1

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and Fig. 8). OsWRKY66 and OsWRKY37 are evolutionary conserved and also expresses

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highly in roots. This indicates that function of OsWRKY66 and OsWRKY37 in root of rice is evolutionary conserved. Similarly, AtWRKY members of GroupIIb are overexpressed in root

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tissues in comparison to other vegetative and floral tissues. On contrary, rice and cotton WRKY members exhibited with moderate to low expression levels amongst studied tissues

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Discussion

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in monocot and dicot plant species.

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(Fig. 8B). These data revealed the diversity in gene expression patterns of WRKY members

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The biological and molecular complexity of an organism certainly intensified in due course of evolution, especially the transcription factors and their binding sites in the upstream regulatory regions of genes have undergone tremendous section pressure. In result, novel regulatory regions/motifs have evolved in the upstream regulatory genic-regions and provide evolutionary platform for functional expansion and diversification (Eulgem et al., 2000). One of such regulatory factors is WRKY transcription factor. WRKY recognizes the conserved DNA motif (T)(T)TGAC(C/T) which is typically known as W-box in the regulatory region. WRKY protein has 60 amino acid long polypeptide chain with 6-amino acid long N-terminal region with highly conserved sequence (WRKYGQ) across species (Rushton et al., 1995). Based on WRKY phylogeny and number of domains present in this protein, WRKY family members are divided in three main groups- Group I, II and III (Fig. 1). Previously, genomewide identification and characterization of WRKY gene sequences amongst various plant species classified all gene members into at least three major groups (Eulgem et al., 2000; Wu et al., 2005; Zhang et al., 2019). The evolutionary comparison of WRKY gene sequences in

Journal Pre-proof the present study also revealed three major groups of gene members and could be further classified into independent sub-groups (Fig. 1). WRKY members of Group I contains two domains in their protein whereas Group II members have only one WRKY domain (Zhang and Wang, 2005). In Group III, WRKY members have one domain but with alerted pattern of zinc finger motif. Group I and II have C2H2 zinc finger pattern whereas in group III WRKY domain contain C2-CH pattern of zinc finger (Eulgem et al., 2000; Rinerson et al., 2015). Group II is further divided into five sub-groups (IIa, IIb, IIc, IId and IIe) based on the gene sequences of WRKY members (Eulgem et al., 2000). This classification is conserved amongst monocot and dicot plant species and is also evident in our results (Fig. 1). Correspondingly,

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WRKY genes have also been grouped into different phylo-groups based on their sequence

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homologies in various plant species including Arabidopsis (Song and Gao, 2014), rice (Wu et al., 2005) and cotton (Dou et al., 2014). Largely, the roles of WRKY trans-factors have been

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characterized as plant transcription factors, however, orthologous sequences of WRKY genes are also present in protozoans such as Dictyostelium discoideum and protozoan-like Giardia

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lamblia (Ulker and Somssich, 2004; Pan et al., 2009; Phukan et al., 2015). Evidently, WRKY genes have also been identified in green algae and in mosses. Though it is believed that

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angiosperm are the descended of green algae, hence WRKY gene sequences were contributed by these ancestors and were further diversified in the plantae kingdom. This suggested that

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WRKY transcription factors are ancient in their origin and also suggest the possibility of lateral gene transfer million years ago (Phukan et al., 2015). Considerably, Group I of

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ancestral WRKY genes is assumed to be the most ancient group as evident by their presence in Chlamydomonas reinhardtii (Fig. 1), and further diversified amongst plant lineages perhaps via genera-specific chromosomal duplication events and also through whole genome merger (polyploidization) events at the evolutionary scale (Ohta, 2000; Zhang and Wang, 2005; Brand et al., 2013). However, it has also been hypothesized that WRKY members of Group I were further individually and parallelly evolved through the domain duplication in protoWRKY gene sequence (Zhang and Wang, 2005). Simultaneously, absence of Group II WRKY gene sequences in spike-moss Selaginella moellendorffii, confirmed its recent divergence at evolutionary scale (Banks et al., 2011; Phukan et al., 2015). Apparently, DNA binding properties of WRKY proteins are not restricted to W-boxes of regulatory regions, because it is evident that OsWRKY13 binds to PRE4 element of 5’-regulatory region of rice genes (Cai et al., 2008); and HvWRKY46 binds with a sugar responsive element (SURE) in barley (Sun et al., 2003).

Journal Pre-proof Previously, evolutionary gene structure and conserved motif analysis revealed the phylogenetic clustering of protein sequences with preserved homologous motif-compositions (Zhang et al., 2019). In the present study, we established the evolutionary trends of 266 paralogs and orthologs members of WRKY gene family. Apparently, divergences in the exon-intron structures of WRKY members were prevalent amongst paralogs and orthologs of WRKY gene family, and the structural divergences in such sequences was accomplished by the exon/intron-gain and insertion/deletion mechanisms at an evolutionary pace (Fig. 4, 5). However, an ever-lasting evolutionary question remains, whether the evolution of gene families in plant system is directed by the exonization of non-coding sequences; or by

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conversion of intronic regions into exonic sequences for the evolution of novel functions, and

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at a faster rate? Exonic length and sequence comparisons revealed that the latter has more possibilities of splitting the existing 175-178 bp long exon into smaller sized exons during

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WRKY gene family evolution (Fig. 5). Henceforth, structural variations amongst the WRKY members exhibited notable ‘degree of nativeness’ in the representative 3D structures (Fig. 7).

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However, paralogs’ sequence alignment showed a degree of conservation in the WRKY

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domains.

Besides the basic cellular reported functions of WRKY gene family, especially in seed

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development, seed dormancy and germination, flowering, trichome and cotton fiber development (Johnson et al., 2002; Luo et al., 2005; Luo et al., 2013), remarkably, Group III

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of WRKY transcription factors involved in response to the infestation of herbivore-, pathogen and nematodes (Song et al., 2018). Correspondingly, AtWRKY54 and AtWRKY70 double mutant of Arabidopsis was tolerant to the osmotic stress (Li et al., 2013). WRKY gene members show large diversity in their gene sequences and subsequent functions also such as AtWRKY50 and AtWRKY51 are involved in the regulation of salicylic pathway and jasmonic acid pathway (Gao et al., 2011); overexpression of BnWRK33 in oilseed rape led to enhanced resistance against Sclerotinia sclerotiorum (Wang et al., 2014b); AtWRKY33 is required for resistance against fungi Alternaria brassicicola and Botrytis cinerea (Zheng et al., 2006); and AtWRKY3 and AtWRKY4 in Arabidopsis

confers resistance against B. cinereal.

Overexpression of AtWRKY41 in Arabidopsis and GhWRKY41 in tobacco enhanced resistant against Pseudomonas and tolerance against salt and drought stresses by regulating stomatal conductance and oxidative stress response (Higashi et al., 2008; Chu et al., 2015), respectively. AtWRKY71 regulates Flowering locus T (FT) and Leafy (LFY) gene thus is involved in regulation of flowering time (Guo et al., 2015). GhWRKY1 in Arabidopsis led to

Journal Pre-proof an increase in the auxin sensitivity but also led to attenuation of Pi starvation stress symptoms (Xu et al., 2012). WRKY transcription factor are also known to regulate production of secondary metabolites like flavanols, tannins (Wang et al., 2010). WRKY in Gossypium arboretum is known to regulate sesquiterpene cyclase, thus enhancing gossypol production (Xu et al., 2004). AaWRKY1 in trichome of Artimisiaannua regulates production of artmisinin (an antimalarial drug) (Ma et al., 2009). During expression analysis of WRKY genes in diverse plant species, we also observed that GhWRKY58 was highly expressed in leaves, root and anther tissues whereas GhWRKY57 were expressed significantly less in all these tissues (Fig. 8). This indicates that GhWRKY57 and GhWRKY58 may have sequence conservation

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largely but are not functionally conserved. Remarkably, the spatial and temporal gene

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expression patterns of WRKY genes in plants update our understanding about their contributions in the regulation of cellular functions as well as their employment into future

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crop improvement programs.

Journal Pre-proof Table 1. dN/dS analysis of WRKY gene family in the representative plant species.

Arabidopsis

dS 0.4393 0.5749 0.5093 0.4807 0.5577 0.651 0.5626 0.5453 0.5453

dN 0.0902 0.1374 0.1159 0.1068 0.1082 0.1511 0.1207 0.1534 0.1534

dN/dS 0.2054 0.2391 0.2277 0.2221 0.194 0.2321 0.2145 0.2813 0.2813

Selection Purifying Purifying Purifying Purifying Purifying Purifying Purifying Purifying Purifying

OsWRKY61 OsWRKY1 OsWRKY15 OsWRKY25 OsWRKY26 OsWRKY40 OsWRKY50 OsWRKY34 OsWRKY38

OsWRKY81 OsWRKY99 OsWRKY98 OsWRKY44 OsWRKY59 OsWRKY64 OsWRKY65 OsWRKY36 OsWRKY53

104.8 267.2 116.8 141.1 135 218.2 233.1 114.8 194.1

498.2 1472.8 705.2 752.9 558 741.8 744.9 623.2 1266.9

0.0305 0.0039 0 0.0945 0.055 0.094 0.1649 0 0

0.0101 0.0014 0 0.0094 0.011 0.0553 0.098 0 0

0.3313 0.3437 0.1721 0.0993 0.2008 0.5888 0.5939 0.001 0.4864

Purifying Purifying Purifying Purifying Purifying Purifying Purifying Purifying Purifying

AtWRKY63 AtWRKY64

AtWRKY64 AtWRKY67

541.2 546.9

0.4012 0.2295

0.1438 0.1855

0.3586 0.8082

Purifying Purifying

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N 1317.7 724.7 932.5 1287 724.8 727 754.6 728.3 728.3

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Rice

S 359.3 277.3 270.5 372.0 205.2 257 229.4 261.7 261.7

na

Cotton

Gene Pairs GhWRKY14 GhWRKY17 GhWRKY12 GhWRKY41 GhWRKY14 GhWRKy22 GhWRKY17 GhWRKY22 GhWRKY30 GhWRKY111 GhWRKY12 GhWRKY69 GhWRKY47 GhWRKY111 GhWRKY41 GhWRKY69 GhWRKY69 GhWRKY84

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Species

181.8 200.1

Journal Pre-proof Acknowledgement This work was supported by the Department of Biotechnology (DBT), Government of India (Grant No. BT/PR8727/AGR/36/777/2013); and Council of Scientific and Industrial Research (CSIR), Government of India (Grant No. 38(1298)/11/EMR-II).

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Conflict of Interest

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The authors declare no conflict of interest.

Journal Pre-proof Legends to the Figures Figure 1. The evolutionary origin of WRKY protein family across the kingdoms. 266 WRKY sequences aligned to construct phylogenetic tree. 265 from plantae (179 dicots and 86 monocots) and 1 sequence from lower plant amino acid sequence aligned by using CLUSTALW and unrooted tree was generated using Interactive Tree of Life (iTOL) tool. The branches of the resulting tree represent individual WRKY sequences and branch lengths are proportionate to the evolutionary distances between and amongst sequences. Nodes represent WRKY sequence divergences occurred by either species divergence or local gene

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duplications (paralog formation). Color indicatives are for domains from different species. Figure 2. Evolutionary history of WRKY domain. The sequence homologies amongst green

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algae, monocots and dicots species were investigated through multiple sequence alignment

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and displayed as WebLogo3 tool plots; and mapped on the evolutionary tree of life. The evolutionary homology patterns were derived by multiple sequence alignment and abstracted

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using WebLogo3 tool plots.

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Figure 3. The evolutionary conserved amino acid residues obtained by using ConSurf server. The conservation score is represented using coloring scheme; conserved amino acids are

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shown in Bordeaux color, average conservation is shown in white color, and variable amino acids are shown in turquoise color.

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Figure 4. Gene structure of eudicot candidates from WRKY gene family. Candidate WRKY gene sequence were compared with their respective cDNA sequences. Yellow boxes represent 5’ to 3’ exonic and black lines indicates intronic regions. The 5’-UTR and 3’UTR are also shown with blue color. The precise length of exon and intron regions (in base pairs) could be estimated from the scale given at the bottom. Figure 5. Determination of exon and intron structural divergence in plantae. The exonic signature sequences of representative eudicot and monocot species were compared and visualized using PIECE online platform. Lengths of exonic and intronic regions can be approximated by scales provided at the top. The colored lines edged by exonic regions signifies matches or mismatches, intron introgressions in exonic regions and frame sharing between exons. Exact matches are outlined by Red line and cyan lines indicates nearly exact match and a common frame shared by exons; and common framed exons with mismatched sequences are connected by light green line.

Journal Pre-proof Figure 6. Evolutionary conservation of WRKY domain in green algae, rice, cotton and Arabidopsis. The figure suggests high conservation in WRKYGQK domain of candidate sequences. Figure 7. Evolutionary structural comparisons of WRKY domain. Surface diagrams of AtWRKY, OsWRKY, GhWRKY and ChrWRKY crystal structures were drawn by using SWISSMODEL server, highlighting the residues in various colored patterns. Figure 8. Expression profiling of WRKY gene family in monocots and eudicot species. The gene expression data was extracted from NCBI, TAIR and RED database. Heatmap

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generation was performed in MeV software package. The temporal variation in the WRKY transcription is expressed using color indicatives as shown on the top of the heatmap; low

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expression is represented by the green color while high expression is represented by the red

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color.

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Legends to the Supplementary Material

S1 Table : List of all WRKY amino acid sequences used in the present study.

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S2 Table : Gene expression data of several WRKY genes in different plant tissues.

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The authors declare no conflict of interest.

Journal Pre-proof Supplementary Table 1: List of all WRKY amino acid sequences used in the present study.

>AtWRKY1 MAEVGKVLASDMELDHSNETKAVDDVVATTDKAEVIPVAVTRTETVVESLESTDCKELEKLVPHTVASQS EVDVASPVSEKAPKVSESSGALSLQSGSEGNSPFIREKVMEDGYNWRKYGQKLVKGNEFVRSYYRCTHPN CKAKKQLERSAGGQVVDTVYFGEHDHPKPLAGAVPINQDKRSDVFTAVSKEKTSGSSVQTLRQTEPPKIH GGLHVSVIPPADDVKTDISQSSRITGDNTHKDYNSPTAKRRKKGGNIELSPVERSTNDSRIVVHTQTLFD IVNDGYRWRKYGQKSVKGSPYPRSYYRCSSPGCPVKKHVERSSHDTKLLITTYEGKHDHDMPPGRVVTHN NMLDSEVDDKEGDANKTPQSSTLQSITKDQHVEDHLRKKTKTNGFEKSLDQGPVLDEKLKEEIKERSDAN KDHAANHAKPEAKSDDKTTVCQEKAVGTLESEEQKPKTEPAQS

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>AtWRKY2 MAGFDENVAVMGEWVPRSPSPGTLFSSAIGEEKSSKRVLERELSLNHGQVIGLEEDTSSNHNKDSSQSNV FRGGLSERIAARAGFNAPRLNTENIRTNTDFSIDSNLRSPCLTISSPGLSPATLLESPVFLSNPLAQPSP TTGKFPFLPGVNGNALSSEKAKDEFFDDIGASFSFHPVSRSSSSFFQGTTEMMSVDYGNYNNRSSSHQSA EEVKPGSENIESSNLYGIETDNQNGQNKTSDVTTNTSLETVDHQEEEEEQRRGDSMAGGAPAEDGYNWRK YGQKLVKGSEYPRSYYKCTNPNCQVKKKVERSREGHITEIIYKGAHNHLKPPPNRRSGMQVDGTEQVEQQ QQQRDSAATWVSCNNTQQQGGSNENNVEEGSTRFEYGNQSGSIQAQTGGQYESGDPVVVVDASSTFSNDE DEDDRGTHGSVSLGYDGGGGGGGGEGDESESKRRKLEAFAAEMSGSTRAIREPRVVVQTTSDVDILDDGY RWRKYGQKVVKGNPNPRSYYKCTAPGCTVRKHVERASHDLKSVITTYEGKHNHDVPAARNSSHGGGGDSG NGNSGGSAAVSHHYHNGHHSEPPRGRFDRQVTTNNQSPFSRPFSFQPHLGPPSGFSFGLGQTGLVNLSMP GLAYGQGKMPGLPHPYMTQPVGMSEAMMQRGMEPKVEPVSDSGQSVYNQIMSRLPQI

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>AtWRKY3 MAEKEEKEPAKLKSSTGGSRPTISLPPRPFGEMFFSGGVGFSPGPMTLVSNLFSDPDEFKSFSQLLAGAM ASPAAAAVAAAAVVATAHHQTPVSSVGGNGGSGGDVDPRFKQSRPTGLMITQPPGMFTVPPGLSPATLLD SPSFFGLFSPLQGTFGMTHQQALAQVTAQAVQGNNVHMQQSQQSEYPSSTQQQQQQQQASLTEIPSFSSA PRSQIRASVQETSQGQRETSEISVSEHRSQPQNADKPADDGYNWRKYGQKQVKGSDFPRSYYKCTHPACP VKKKVERSLDGQVTEIIYKGQHNHELPQKRGNNNGSCKSSDIANQFQTSNSSLNKSKRDQETSQVTTTEQ MSEASDSEEVGNAETSVGERHEDEPDPKRRNTEVRVSEPVASSHRTVTEPRIIVQTTSEVDLLDDGYRWR KYGQKVVKGNPYPRSYYKCTTPDCGVRKHVERAATDPKAVVTTYEGKHNHDVPAARTSSHQLRPNNQHNT STVNFNHQQPVARLRLKEEQIT >AtWRKY4 MSEKEEAPSTSKSTGAPSRPTLSLPPRPFSEMFFNGGVGFSPGPMTLVSNMFPDSDEFRSFSQLLAGAMS SPATAAAAAATASDYQRLGEGTNSSSGDVDPRFKQNRPTGLMISQSQSPSMFTVPPGLSPAMLLDSPSFL GLFSPVQGSYGMTHQQALAQVTAQAVQANANMQPQTEYPPPSQVQSFSSGQAQIPTSAPLPAQRETSDVT IIEHRSQQPLNVDKPADDGYNWRKYGQKQVKGSEFPRSYYKCTNPGCPVKKKVERSLDGQVTEIIYKGQH NHEPPQNTKRGNKDNTANINGSSINNNRGSSELGASQFQTNSSNKTKREQHEAVSQATTTEHLSEASDGE EVGNGETDVREKDENEPDPKRRSTEVRISEPAPAASHRTVTEPRIIVQTTSEVDLLDDGYRWRKYGQKVV KGNPYPRSYYKCTTPGCGVRKHVERAATDPKAVVTTYEGKHNHDLPAAKSSSHAAAAAQLRPDNRPGGLA NLNQQQQQQPVARLRLKEEQTT >AtWRKY6 MDRGWSGLTLDSSSLDLLNPNRISHKNHRRFSNPLAMSRIDEEDDQKTRISTNGSEFRFPVSLSGIRDRE DEDFSSGVAGDNDREVPGEVDFFSDKKSRVCREDDEGFRVKKEEQDDRTDVNTGLNLRTTGNTKSDESMI DDGESSEMEDKRAKNELVKLQDELKKMTMDNQKLRELLTQVSNSYTSLQMHLVSLMQQQQQQNNKVIEAA EKPEETIVPRQFIDLGPTRAVGEAEDVSNSSSEDRTRSGGSSAAERRSNGKRLGREESPETESNKIQKVN STTPTTFDQTAEATMRKARVSVRARSEAPMISDGCQWRKYGQKMAKGNPCPRAYYRCTMATGCPVRKQVQ RCAEDRSILITTYEGNHNHPLPPAAVAMASTTTAAANMLLSGSMSSHDGMMNPTNLLARAVLPCSTSMAT ISASAPFPTVTLDLTHSPPPPNSSNPSSSVATNNNHNSLMQRPQQQQQQMTNLPPGMLPHVIGQALYNQS KFSGLQFSGGSPSTAAFSQSHAVADTITALTADPNFTAALAAVISSMINGTNHHDGEGNNKNQ

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>AtWRKY7 MTVELMMSSYSGGGGGGDGFPAIAAAAKMEDTALREAASAGIHGVEEFLKLIGQSQQPTEKSQTEITAVT DVAVNSFKKVISLLGRSRTGHARFRRAPASTQTPFKQTPVVEEEVEVEEKKPETSSVLTKQKTEQYHGGG SAFRVYCPTPIHRRPPLSHNNNNNQNQTKNGSSSSSPPMLANGAPSTINFAPSPPVSATNSFMSSHRCDT DSTHMSSGFEFTNPSQLSGSRGKPPLSSASLKRRCNSSPSSRCHCSKKRKSRVKRVIRVPAVSSKMADIP SDEFSWRKYGQKPIKGSPHPRGYYKCSSVRGCPARKHVERALDDAMMLIVTYEGDHNHALVLETTTMNHD KTL

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>AtWRKY8 MSHEIKDLNNYHYTSSYNHYNINNQNMINLPYVSGPSAYNANMISSSQVGFDLPSKNLSPQGAFELGFEL SPSSSDFFNPSLDQENGLYNAYNYNSSQKSHEVVGDGCATIKSEVRVSASPSSSEADHHPGEDSGKIRKK REVGDGGEDDQRSQKVVKTKKKEEKKKEPRVSFMTKTEVDHLEDGYRWRKYGQKAVKNSPYPRSYYRCTT QKCNVKKRVERSYQDPTVVITTYESQHNHPIPTNRRTAMFSGTTASDYNPSSSPIFSDLIINTPRSFSND DLFRVPYASVNVNPSYHQQQHGFHQQESEFELLKEMFPSVFFKQEP

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>AtWRKY9 MGFDFSTSKSKAKRQKRIEVRFASPLMGIDLSLKLEAEEKKKEIEGSKHSRENKEDEEHDASGDEDEQMV KEDEDDSSSLGLRTREEENEREELLQLQIQMESVKEENTRLRKLVEQTLEDYRHLEMKFPVIDKTKKMDL EMFLGVQGKRCVDITSKARKRGAERSPSMEREIGLSLSLEKKQKQEESKEAVQSHHQRYNSSSLDMNMPR IISSSQGNRKARVSVRARCETATMNDGCQWRKYGQKTAKGNPCPRAYYRCTVAPGCPVRKQVQRCLEDMS ILITTYEGTHNHPLPVGATAMASTASTSPFLLLDSSDNLSHPSYYQTPQAIDSSLITYPQNSSYNNRTIR SLNFDGPSRGDHVSSSQNRLNWMM

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>AtWRKY10 MSDFDENFIEMTSYWAPPSSPSPKTILAMLEQTDNGLDPISEIFPQESLPRDHTDQSGQRSGLRERLAAR VGFNLPTLNTEENMSPLDAFFRSSNVPNSPVVAISPGFSPSALLHTPNMVSDSSQIIPPSSATNYGPLEM VETSGEDNAAMMMFNNDLPYQPYNVDLPSLEVFDDIATEESFYIPSYEPDVDPIGTPLVASFESDLIDDA HTDIISIEDSESEDGNKDDDDEDFQYEDEDEDQDQDQDQDVDEDEEEEKDEDNVALDDPRPPSPKRRRYE VSNMIGATRTSKTQRIILQMESDEDNPNDGYRWRKYGQKVVKGNPNPRSYFKCTNIECRVKKHVERGADN IKLVVTTYDGIHNHPSPPARRSNSSSRNRSAGATIPQNQNDRTSRLGRAPPTPPPPPPSSYTPEEMRPFS SLATEIDLTEVYMTGISMLPNIPVYENSGFMDQNSGFMYQNDEPTMNAMPDGSDVYDGIMERLYFKFGVD M >AtWRKY11 MAVDLMRFPKIDDQTAIQEAASQGLQSMEHLIRVLSNRPEQQHNVDCSEITDFTVSKFKTVISLLNRTGH ARFRRGPVHSTSSAASQKLQSQIVKNTQPEAPIVRTTTNHPQIVPPPSSVTLDFSKPSIFGTKAKSAELE FSKENFSVSLNSSFMSSAITGDGSVSNGKIFLASAPLQPVNSSGKPPLAGHPYRKRCLEHEHSESFSGKV SGSAYGKCHCKKRKNRMKRTVRVPAISAKIADIPPDEYSWRKYGQKPIKGSPHPRGYYKCSTFRGCPARK HVERALDDPAMLIVTYEGEHRHNQSAMQENISSSGINDLVFASA >AtWRKY12 MEGGGRRVFSNYDLQQVTSSSTTIQENMNFLVPFEETNVLTFFSSSSSSSLSSPSFPIHNSSSTTTTHAP LGFSNNLQGGGPLGSKVVNDDQENFRGGTNNDAHSNSWWRSNSGSGDMKNKVKIRRKLREPRFCFQTKSD VDVLDDGYKWRKYGQKVVKNSLHPRSYYRCTHNNCRVKKRVERLSEDCRMVITTYEGRHNHIPSDDSTSP DHDCLSSF >AtWRKY13 MGAINQGISLFDESQTVINPINTNHLGFFFSFPSHSTLSSSSSSSSSSPSSLVSPFLGHNSLNSFLHNNP SSFISHPQDSINLMTNLPETLISSLSSSKQRDDHDGFLNLDHHRLTGSISSQRPLSNPWAWSCQAGYGSS QKNNHGSEIDVDDNDDEVGDGGGINDDDNGRHHHHDTPSRHDKHNTASLGVVSSLKMKKLKTRRKVREPR FCFKTLSEVDVLDDGYRWRKYGQKVVKNTQHPRSYYRCTQDKCRVKKRVERLADDPRMVITTYEGRHLHS PSNHLDDDSLSTSHLHPPLSNFFW >AtWRKY14

Journal Pre-proof MCSVSELLDMENFQGDLTDVVRGIGGHVLSPETPPSNIWPLPLSHPTPSPSDLNINPFGDPFVSMDDPLL QELNSITNSGYFSTVGDNNNNIHNNNGFLVPKVFEEDHIKSQCSIFPRIRISHSNIIHDSSPCNSPAMSA HVVAAAAAASPRGIINVDTNSPRNCLLVDGTTFSSQIQISSPRNLGLKRRKSQAKKVVCIPAPAAMNSRS SGEVVPSDLWAWRKYGQKPIKGSPFPRGYYRCSSSKGCSARKQVERSRTDPNMLVITYTSEHNHPWPIQR NALAGSTRSSTSSSSNPNPSKPSTANVNSSSIGSQNTIYLPSSTTPPPTLSSSAIKDERGDDMELENVDD DDDNQIAPYRPELHDHQHQPDDFFADLEELEGDSLSMLLSHGCGGDGKDKTTASDGISNFFGWSGDNNYN NYDDQDSRSL >AtWRKY15 MAVELMTRNYISGVGADSFAVQEAAASGLKSIENFIGLMSRDSFNSDQPSSSSASASASAAADLESARNT TADAAVSKFKRVISLLDRTRTGHARFRRAPVHVISPVLLQEEPKTTPFQSPLPPPPQMIRKGSFSSSMKT IDFSSLSSVTTESDNQKKIHHHQRPSETAPFASQTQSLSTTVSSFSKSTKRKCNSENLLTGKCASASSSG RCHCSKKRKIKQRRIIRVPAISAKMSDVPPDDYSWRKYGQKPIKGSPHPRGYYKCSSVRGCPARKHVERA ADDSSMLIVTYEGDHNHSLSAADLAGAAVADLILESS

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>AtWRKY16 MERIGIYSKLLEIEKMVCKQLLDIRCVGIWGMPGIGKTTLAKAVFDQMSGEFDAHCFIEDYTKAIQEKGV YCLLEEQFLKQNAGASGTVTKLSLLRDRLNNKRVLVVLDDVRSPLVVESFLGGFDWFGPKSLIIITSKDK SVFRLCRVNQIYEVQGLNEKEALQLFSLCASIDDMAEQNLHEVSMKVIKYANGHPLALNLYGRELMGKKR PPEMEIAFLKLKECPPAIFVDAIKSSYDTLNDREKNIFLDIACFFQGENVDYVMQLLEGCGFFPHVGIDV LVEKSLVTISENRVRMHNLIQDVGRQIINRETRQTKRRSRLWEPCSIKYLLEDKEQNENDEQKTTFERAQ VPEEIEGIFLDTSNLSFDIKHVAFDNMLNLRLFKIYSSNPEVHHVNNFLKGSLSSLPNVLRLLHWENYPL QFLPQNFDPIHLVEINMPYSQLKKLWGGTKDLEMLKTIRLCHSQQLVDIDDLLKAQNLEVVDLQGCTRLQ SFPATGQLLHLRVVNLSGCTEIKSFPEIPPNIETLNLQGTGIIELPLSIVKPNYRELLNLLAEIPGLSGV SNLEQSDLKPLTSLMKISTSYQNPGKLSCLELNDCSRLRSLPNMVNLELLKALDLSGCSELETIQGFPRN LKELYLVGTAVRQVPQLPQSLEFFNAHGCVSLKSIRLDFKKLPVHYTFSNCSDLSPQVVNDFLVQAMANV IAKHIPRERHVTGFSQKTVQRSSRDSQQELNKTLAFSFCAPSHANQNSKLDLQPGSSSMTRLDPSWRNTL VGFAMLVQVAFSEGYCDDTDFGISCVCKWKNKEGHSHRREINLHCWALGKAVERDHTFVFFDVNMRPDTD EGNDPDIWADLVVFEFFPVNKQRKPLNDSCTVTRCGVRLITAVNCNTSIENISPVLSLDPMEVSGNEDEE VLRVRYAGLQEIYKALFLYIAGLFNDEDVGLVAPLIANIIDMDVSYGLKVLAYRSLIRVSSNGEIVMHYL LRQMGKEILHTESKKTDKLVDNIQSSMIATKEIEITRSKSRRKNNKEKRVVCVVDRGSRSSDLWVWRKYG QKPIKSSPYPRSYYRCASSKGCFARKQVERSRTDPNVLVITYISEHNHPFPTLRNTLAGSTRSSSSKCSD VTTSASSTVSQDKEGPDKSHLPSSPASPPYAAMVVKEEDMEQWDNMEFDVDVEEDTFIPELFPEDTFADM DKLEENSQTMFLSRRSSGGNMEAQGKNSSDDREVNLPSKILNR >AtWRKY17 MTVDIMRLPKMEDQTAIQEAASQGLKSMEHLIRVLSNRPEERNVDCSEITDFTVSKFKKVISLLNRSGHA RFRRGPVHSPPSSSVPPPVKVTTPAPTQISAPAPVSFVQANQQSVTLDFTRPSVFGAKSKSSEVVEFAKE SFSVSSNSSFMSSAITGDGSVSKGSSIFLAPAPAVPVTSSGKPPLSGLPYRKRCFEHDHSEGFSGKISGS GNGKCHCKKSRKNRMKRTVRVPAVSAKIADIPPDEYSWRKYGQKPIKGSPHPRGYYKCSTFRGCPARKHV ERALDDSTMLIVTYEGEHRHHQSTMQENITPSVSGLVFGSA >AtWRKY18 MDGSSFLDISLDLNTNPFSAKLPKKEVSVLASTHLKRKWLEQDESASELREELNRVNSENKKLTEMLARV CESYNELHNHLEKLQSRQSPEIEQTDIPIKKRKQDPDEFLGFPIGLSSGKTENSSSNEDHHHHHQQHEQK NQLLSCKRPVTDSFNKAKVSTVYVPTETSDTSLTVKDGFQWRKYGQKVTRDNPSPRAYFRCSFAPSCPVK KKVQRSAEDPSLLVATYEGTHNHLGPNASEGDATSQGGSSTVTLDLVNGCHRLALEKNERNNTMQEVLIQ QMASSLTKDSKFTAALAAAISGRLMEQSRT >AtWRKY19 MAATPPISSGSGLSQQLNNSSSSKLCQVEGCQKGARDASGRCISHGGGRRCQKPDCQKGAEGKTVYCKAH GGGRRCEYLGCTKGAEGSTDFCIAHGGGRRCNHEDCTRSAWGRTEFCVKHGGGARCKTYGCGKSASGPLP FCRAHGGGKKCSHEDCTGFARGRSGLCLMHGGGKRCQRENCTKSAEGLSGLCISHGGGRRCQSIGCTKGA KGSKMFCKACITKRPLTIDGGGNMGGVTTGDALNYLKAVKDKFEDSEKYDTFLEVLNDCKHQGVDTSGVI ARLKDLFKGHDDLLLGFNTYLSKEYQITILPEDDFPIDFLDKVEGPYEMTYQQAQTVQANANMQPQTEYP SSSAVQSFSSGQPQIPTSAPDSSLLAKSNTSGITIIEHMSQQPLNVDKQVNDGYNWQKYGQKKVKGSKFP LSYYKCTYLGCPSKKKVERSLDGQVAEIVYKDRHNHEPPNQGKDGSTTYLSGSSTHINCMSSELTASQFS

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SNKTKIEQQEATSLATTIEYMSEASDNEEDSNGETSEGEKDEDEPEPKRRITEVQVSELADASDRTVREP RVIFQTTSEVDNLDDGYRWRKYGQKVVKGNPYPRFSSSKDYDVVIRYGRADISNEDFISHLRASLCRRGI SVYEEFNEVDALPKCRVLIIVLTSTYVPSNLLNILEHQHTEDRVVYPIFYRLSPYDFVCNSKNYERFYLQ DEPKKWQAALKEITQMPGYTLTDKSESELIDEIVRDALKVLCSADKVNMIGMDMQVEEILSLLCIESLDV RSIGIWGTVGIGKTTIAEEIFRKISVQYETCVVLKDLHKEVEVKGHDAVRENFLSEVLEVEPHVIRISDI KTSFLRSRLQRKRILVILDDVNDYRDVDTFLGTLNYFGPGSRIIMTSRNRRVFVLCKIDHVYEVKPLDIP KSLLLLDRGTCQIVLSPEVYKTLSLELVKFSNGNPQVLQFLSSIDREWNKLSQEVKTTSPIYIPGIFEKS CCGLDDNERGIFLDIACFFNRIDKDNVAMLLDGCGFSAHVGFRGLVDKSLLTISQHNLVDMLSFIQATGR EIVRQESADRPGDRSRLWNADYIRHVFINDTGTSAIEGIFLDMLNLKFDANPNVFEKMCNLRLLKLYCSK AEEKHGVSFPQGLEYLPSKLRLLHWEYYPLSSLPKSFNPENLVELNLPSSCAKKLWKGKKSLEKLKKMRL SYSDQLTKIPRLSSATNLEHIDLEGCNSLLSLSQSISYLKKLVFLNLKGCSKLENIPSMVDLESLEVLNL SGCSKLGNFPEISPNVKELYMGGTMIQEIPSSIKNLVLLEKLDLENSRHLKNLPTSIYKLKHLETLNLSG CISLERFPDSSRRMKCLRFLDLSRTDIKELPSSISYLTALDELLFVDSRRNSPVVTNPNANSTELMPSES SKLEILGTPADNEVVVGGTVEKTRGIERTPTILVKSREYLIPDDVVAVGGDIKGLRPPVLQLQPAMKLSH IPRGSTWDFVTHFAPPETVAPPSSSSEAREEEVETEETGAMFIPLGDKETCSFTVNKGDSSRTISNTSPI YASEGSFITCWQKGQLLGRGSLGSVYEGISADGDFFAFKEVSLLDQGSQAHEWIQQVEGGIALLSQLQHQ NIVRYRGTTKDESNLYIFLELVTQGSLRKLYQRNQLGDSVVSLYTRQILDGLKYLHDKGFIHRNIKCANV LVDANGTVKLADFGLAKVMSLWRTPYWNWMAPEVIVLKSFPLF

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>AtWRKY20 MNPQANDRKEFQGDCSATGDLTAKHDSAGGNGGGGARYKLMSPAKLPISRSTDITIPPGLSPTSFLESPV FISNIKPEPSPTTGSLFKPRPVHISASSSSYTGRGFHQNTFTEQKSSEFEFRPPASNMVYAELGKIRSEP PVHFQGQGHGSSHSPSSISDAAGSSSELSRPTPPCQMTPTSSDIPAGSDQEESIQTSQNDSRGSTPSILA DDGYNWRKYGQKHVKGSEFPRSYYKCTHPNCEVKKLFERSHDGQITDIIYKGTHDHPKPQPGRRNSGGMA AQEERLDKYPSSTGRDEKGSGVYNLSNPNEQTGNPEVPPISASDDGGEAAASNRNKDEPDEDDPFSKRRR MEGAMEITPLVKPIREPRVVVQTLSEVDILDDGYRWRKYGQKVVRGNPNPRSYYKCTAHGCPVRKHVERA SHDPKAVITTYEGKHDHDVPTSKSSSNHEIQPRFRPDETDTISLNLGVGISSDGPNHASNEHQHQNQQLV NQTHPNGVNFRFVHASPMSSYYASLNSGMNQYGQRETKNETQNGDISSLNNSSYPYPPNMGRVQSGP

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>AtWRKY21 MEEIEGTNRAAVESCHRVLNLLHRSQQQDHVGFEKNLVSETREAVIRFKRVGSLLSSSVGHARFRRAKKL QSHVSQSLLLDPCQQRTTEVPSSSSQKTPVLRSGFQELSLRQPSDSLTLGTRSFSLNSNAKAPLLQLNQQ TMPPSNYPTLFPVQQQQQQQQQQQQQQQQFHERLQAHHLHQQQQLQKHQAELMLRKCNGGISLSFDNSSC TPTMSSTRSFVSSLSIDGSVANIEGKNSFHFGVPSSTDQNSLHSKRKCPLKGDEHGSLKCGSSSRCHCAK KRKHRVRRSIRVPAISNKVADIPPDDYSWRKYGQKPIKGSPYPRGYYKCSSMRGCPARKHVERCLEDPAM LIVTYEAEHNHPKLPSQAITT >AtWRKY22 MADDWDLHAVVRGCSAVSSSATTTVYSPGVSSHTNPIFTVGRQSNAVSFGEIRDLYTPFTQESVVSSFSC INYPEEPRKPQNQKRPLSLSASSGSVTSKPSGSNTSRSKRRKIQHKKVCHVAAEALNSDVWAWRKYGQKP IKGSPYPRGYYRCSTSKGCLARKQVERNRSDPKMFIVTYTAEHNHPAPTHRNSLAGSTRQKPSDQQTSKS PTTTIATYSSSPVTSADEFVLPVEDHLAVGDLDGEEDLLSLSDTVVSDDFFDGLEEFAAGDSFSGNSAPA SFDLSWVVNSAATTTGGI >AtWRKY23 MEFTDFSKTSFYYPSSQSVWDFGDLAAAERHSLGFMELLSSQQHQDFATVSPHSFLLQTSQPQTQTQPSA KLSSSIIQAPPSEQLVTSKVESLCSDHLLINPPATPNSSSISSASSEALNEEKPKTEDNEEEGGEDQQEK SHTKKQLKAKKNNQKRQREARVAFMTKSEVDHLEDGYRWRKYGQKAVKNSPFPRSYYRCTTASCNVKKRV ERSFRDPSTVVTTYEGQHTHISPLTSRPISTGGFFGSSGAASSLGNGCFGFPIDGSTLISPQFQQLVQYH >AtWRKY24 MDREDINPMFSRLDVENNNTFSSFVDKTLMMMPPSTFSGEVEPSSSSSWYPESFHVHAPPLPPENDQIGE KGKELKEKRSRKVPRIAFHTRSDDDVLDDGYRWRKYGQKSVKHNAHPRSYYRCTYHTCNVKKQVQRLAKD PNVVVTTYEGVHNHPCEKLMETLNPLLRQLQFLSSFSNL >AtWRKY25 MSSTSFTDLLGSSGVDCYEDDEDLRVSGSSFGGYYPERTGSGLPKFKTAQPPPLPISQSSHNFTFSDYLD SPLLLSSSHSLISPTTGTFPLQGFNGTTNNHLDFPWQLQSQPSNASSALQETYGVQDHEKKQEMIPNEIE TQNNNQSFGTERQIKIPAYMVSRNSNDGYGWRKYGQKQVKKSENPRSYFKCTYPDCVSKKIVETASDGQI

Journal Pre-proof TEIIYKGGHNHPKPEFTKRPSQSSLPSSVNGRRLFNPASVVSEPHDQSENSSISFDYSDLEQKSFKSEYG EIDEEEEQPEMKRMKREGEDEGMSIEVSKGVKEPRVVVQTISDIDVLIDGFRWRKYGQKVVKGNTNPRSY YKCTFQGCGVKKQVERSAADERAVLTTYEGRHNHDIPTALRRS >AtWRKY26 MVTQPLPQLDLFKSEIMSSNKTSDDGYNWRKYGQKQVKGSENPRSYFKCTYPNCLAKKKEETSLVKGQMI EIVYKGSHNHPKPQSTKRSSSTAIAAHQNSSNGDGKDIGEDETEAKRWKREENVKEPRVVVQTTSDIDIL DDGYRWRKYGQKVVKGNPNPRSYYKCTFTGCFVRKHVERAFQDPKSVITTYEGKHKHQIPTPRRGPVLRL LGKTET

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>AtWRKY27 MSSEDWDLFAVVRSCSSSVSTTNSCAGHEDDIGNCKQQQDPPPPPLFQASSSCNELQDSCKPFLPVTTTT TTTWSPPPLLPPPKASSPSPNILLKQEQVLLESQDQKPPLSVRVFPPSTSSSSVFVFRGQRDQLLQQQSQ PPLRSRKRKNQQKRTICHVTQENLSSDLWAWRKYGQKPIKGSPYPRNYYRCSSSKGCLARKQVERSNLDP NIFIVTYTGEHTHPRPTHRNSLAGSTRNKSQPVNPVPKPDTSPLSDTVKEEIHLSPTTPLKGNDDVQETN GDEDMVGQEVNMEEEEEEEEVEEDDEEEEDDDDVDDLLIPNLAVRDRDDLFFAGSFPSWSAGSAGDGGG

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>AtWRKY28 MSNETRDLYNYQYPSSFSLHEMMNLPTSNPSSYGNLPSQNGFNPSTYSFTDCLQSSPAAYESLLQKTFGL SPSSSEVFNSSIDQEPNRDVTNDVINGGACNETETRVSPSNSSSSEADHPGEDSGKSRRKRELVGEEDQI SKKVGKTKKTEVKKQREPRVSFMTKSEVDHLEDGYRWRKYGQKAVKNSPYPRSYYRCTTQKCNVKKRVER SFQDPTVVITTYEGQHNHPIPTNLRGSSAAAAMFSADLMTPRSFAHDMFRTAAYTNGGSVAAALDYGYGQ SGYGSVNSNPSSHQVYHQGGEYELLREIFPSIFFKQEP

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>AtWRKY29 MGEVAYMDEGDLEAIVRGYSGSGDAFSGESSGTFSPSFCLPMETSSFYEPEMETSGLDELGELYKPFYPF STQTILTSSVSLPEDSKPFRDDKKQRSHGCLLSNGSRADHIRISESKSKKSKKNQQKRVVEQVKEENLLS DAWAWRKYGQKPIKGSPYPRSYYRCSSSKGCLARKQVERNPQNPEKFTITYTNEHNHELPTRRNSLAGST RAKTSQPKPTLTKKSEKEVVSSPTSNPMIPSADESSVAVQEMSVAETSTHQAAGAIEGRRLSNGLPSDLM SGSGTFPSFTGDFDELLNSQEFFSGYLWNY

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>AtWRKY30 MEKNHSSGEWEKMKNEINELMIEGRDYAHQFGSASSQETREHLAKKILQSYHKSLTIMNYSGELDQVSQG GGSPKSDDSDQEPLVIKSSKKSMPRWSSKVRIAPGAGVDRTLDDGFSWRKYGQKDILGAKFPRGYYRCTY RKSQGCEATKQVQRSDENQMLLEISYRGIHSCSQAANVGTTMPIQNLEPNQTQEHGNLDMVKESVDNYNH QAHLHHNLHYPLSSTPNLENNNAYMLQMRDQNIEYFGSTSFSSDLGTSINYNFPASGSASHSASNSPSTV PLESPFESYDPNHPYGGFGGFYS

>AtWRKY31 MFRFPVSLGGSRDEDRHDQITPLDDHRVVVDEVDFFSEKRDRVSRENINDDDDEGNKVLIKMEGSRVEEN DRSRDVNIGLNLLTANTGSDESTVDDGLSMDMEDKRAKIENAQLQEELKKMKIENQRLRDMLSQATTNFN ALQMQLVAVMRQQEQRNSSQDHLLAQESKAEGRKRQELQIMVPRQFMDLGPSSGAAEHGAEVSSEERTTV RSGSPPSLLESSNPRENGKRLLGREESSEESESNAWGNPNKVPKHNPSSSNSNGNRNGNVIDQSAAEATM RKARVSVRARSEAAMISDGCQWRKYGQKMAKGNPCPRAYYRCTMAGGCPVRKQVQRCAEDRSILITTYEG NHNHPLPPAAMAMASTTTAAASMLLSGSMSSQDGLMNPTNLLARAILPCSSSMATISASAPFPTITLDLT NSPNGNNPNMTTNNPLMQFAQRPGFNPAVLPQVVGQAMYNNQQQSKFSGLQLPAQPLQIAATSSVAESVS AASAAIASDPNFAAALAAAITSIMNGSSHQNNNTNNNNVATSNNDSRQ >AtWRKY32 MEEDTGIDEAKTYTVEKSEKVEPEKDGLSQFRDEEKSLGADMEDLHDETVRETLGKDQVQGVRENSSVEP NVEDVLEVNETDSVKETVVSAIVPVDEVEENRQVETSPSLAASSDSLTVTPCRSLDPATASTAQDLPLVS VPTKQEQRSDSPVVNRLSVTPVPRTPARDGYNWRKYGQKQVKSPKGSRSYYRCTYTECCAKKIECSNDSG NVVEIVNKGLHTHEPPRKTSFSPREIRVTTAIRPVSEDDTVVEELSIVPSGSDPSASTKEYICESQTLVD

Journal Pre-proof RKRHCENEAVEEPEPKRRLKKDNSQSSDSVSKPGKKNKFVVHAAGDVGICGDGYRWRKYGQKMVKGNPHP RNYYRCTSAGCPVRKHIETAVENTKAVIITYKGVHNHDMPVPKKRHGPPSSMLVAAAAPTSMRTRTDDQV NIPTSSQCSVGRESEKQSKEALDVGGEKVMESARTLLSIGFEIKQC >AtWRKY33 MAASFLTMDNSRTRQNMNGSANWSQQSGRTSTSSLEDLEIPKFRSFAPSSISISPSLVSPSTCFSPSLFL DSPAFVSSSANVLASPTTGALITNVTNQKGINEGDKSNNNNFNLFDFSFHTQSSGVSAPTTTTTTTTTTT TTNSSIFQSQEQQKKNQSEQWSQTETRPNNQAVSYNGREQRKGEDGYNWRKYGQKQVKGSENPRSYYKCT FPNCPTKKKVERSLEGQITEIVYKGSHNHPKPQSTRRSSSSSSTFHSAVYNASLDHNRQASSDQPNSNNS FHQSDSFGMQQEDNTTSDSVGDDEFEQGSSIVSRDEEDCGSEPEAKRWKGDNETNGGNGGGSKTVREPRI VVQTTSDIDILDDGYRWRKYGQKVVKGNPNPRSYYKCTTIGCPVRKHVERASHDMRAVITTYEGKHNHDV PAARGSGYATNRAPQDSSSVPIRPAAIAGHSNYTTSSQAPYTLQMLHNNNTNTGPFGYAMNNNNNNSNLQ TQQNFVGGGFSRAKEEPNEETSFFDSFMP

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>AtWRKY34 MAGIDNKAAVMGEWFDCSTTNHRKRSKAELGREFSLNYIKNEDSLQITFQESSRGALRERIAARSGFNAP WLNTEDILQSKSLTISSPGLSPATLLESPVFLSNPLLSPTTGKLSSVPSDKAKAELFDDITTSLAFQTIS GSGLDPTNIALEPDDSQDYEERQLGGLGDSMACCAPADDGYNWRKYGQKLVKGSEYPRSYYKCTHPNCEA KKKVERSREGHIIEIIYTGDHIHSKPPPNRRSGIGSSGTGQDMQIDATEYEGFAGTNENIEWTSPVSAEL EYGSHSGSMQVQNGTHQFGYGDAAADALYRDENEDDRTSHMSVSLTYDGEVEESESKRRKLEAYATETSG STRASREPRVVVQTTSDIDILDDGYRWRKYGQKVVKGNPNPRSYYKCTANGCTVTKHVERASDDFKSVLT TYIGKHTHVVPAARNSSQVGAGSSGTLQGSLATQTHNHNVHYPMPHSRSEGLATANSSLFDFQSHLRHPT GFSVYIGQSELSDLSMPGLTIGQEKLTSLQAPDIGDPTGLMLQLAAQPKVEPVSPQQGLDLSASSLICRE MLSRLRQI

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>AtWRKY35 MDNFQGDLTDVVRGIGSGHVSPSPGPPEGPSPSSMSPPPTSDLHVEFPSAATSASCLANPFGDPFVSMKD PLIHLPASYISGAGDNKSNKSFAIFPKIFEDDHIKSQCSVFPRIKISQSNNIHDASTCNSPAITVSSAAV AASPWGMINVNTTNSPRNCLLVDNNNNTSSCSQVQISSSPRNLGIKRRKSQAKKVVCIPAPAAMNSRSSG EVVPSDLWAWRKYGQKPIKGSPYPRGYYRCSSSKGCSARKQVERSRTDPNMLVITYTSEHNHPWPTQRNA LAGSTRSSSSSSLNPSSKSSTAAATTSPSSRVFQNNSSKDEPNNSNLPSSSTHPPFDAAAIKEENVEERQ EKMEFDYNDVENTYRPELLQEFQHQPEDFFADLDELEGDSLTMLLSHSSGGGNMENKTTIPDVFSDFFDD DESSRSL >AtWRKY36 MIKEETVSYFQTFDGVMAESDKEEELDATKAKVEKVREENEKLKLLLSTILNNYNSLQMQVSKVLGQQQG ASSMELDHIDRQDENNDYDVDISLRLGRSEQKISKKEENKVDKISTKDVEESKDKRSALGFGFQIQSYEA SKLDDLCRQVKLANAENKCVSSRKDVKTVRNENHQDVLEEHEQTGLKKTRVCVKASCEDPSINDGCQWRK YGQKTAKTNPLPRAYYRCSMSSNCPVRKQVQRCGEEETSAFMTTYEGNHDHPLPMEASHMAAGTSAAASL LQSGSSSSSSSTSASLSYFFPFHHFSISTTNSHPTVTLDLTRPNYPNQLPDDYPLSSSSFSLNFSSPDPP PPSSHDHTLNFSGLRTQAPLSTDSLLARYRTRLSGQQ >AtWRKY38 MEMNSPHEKAVQAIRYGHSCAMRLKRRLNHPMADGGPLSSYDLAKSIVESFSNAISILSAKPETEDDQFS DLSSRDSSPPPQGSPSKKRKIDSTNSSENWRDDSPDPIYYDGYLWRKYGQKSIKKSNHQRSYYRCSYNKD HNCEARKHEQKIKDNPPVYRTTYFGHHTCKTEHNLDAIFIAGQDPLDDFKSTQMIRFGKDQDQEKESRSN GFSLSVKHEEDIIKEQAIDQYREITSNDQDCQDVIEEYLSSPSGSYPPSSSSGSESADFNSDLLFDNPDS WDRYDQFYF >AtWRKY39 MEEVEAANRSAIESCHGVLNLLSQRTSDPKSITVETGEVVSKFKRVASLLTRGLGHGKFRSTNKFRSSFP QHIFLESPICCGNDLSGDYTQVLAPEPLQMVPASAVYNEMEPKHQLGHPSLMLSHKMCVDKSFLELKPPP FRAPYQLIHNHQQIAYSRSNSGVNLKFDGSGSSCYTPSVSNGSRSFVSSLSMDASVTDYDRNSFHLTGLS RGSDQQHTRKMCSGSLKCGSRSKCHCSKKRKLRVKRSIKVPAISNKIADIPPDEYSWRKYGQKPIKGSPH PRGYYKCSSVRGCPARKHVERCIDETSMLIVTYEGEHNHSRILSSQSAHT

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>AtWRKY40 MDQYSSSLVDTSLDLTIGVTRMRVEEDPPTSALVEELNRVSAENKKLSEMLTLMCDNYNVLRKQLMEYVN KSNITERDQISPPKKRKSPAREDAFSCAVIGGVSESSSTDQDEYLCKKQREETVVKEKVSRVYYKTEASD TTLVVKDGYQWRKYGQKVTRDNPSPRAYFKCACAPSCSVKKKVQRSVEDQSVLVATYEGEHNHPMPSQID SNNGLNRHISHGGSASTPVAANRRSSLTVPVTTVDMIESKKVTSPTSRIDFPQVQKLLVEQMASSLTKDP NFTAALAAAVTGKLYQQNHTEK >AtWRKY41 MEIMNWERRSLLNELIHGLNAAKQLQGSSSPSLTASSSCLTTEIKETLLHNIVSSFKKAILMLNGSTTQH NPTIELAPDHLAHPGKVSESPASIIGSPRSEEFFDVGSKEFNLSSKKRKMLPKWTEQVRISPERGLEGPH DDIFSWRKYGQKDILGAKFPRSYYRCTYRNIQYCWATKQVQRSDGDPTIFEVTYRGTHTCSQGIPPPEKR ETKPNHTVVVNYQNNLLDNLRMVRTGRLGSEAFSFPVRSPLYTYESINGGGTFYHHVGSSGPSDFTGLIS TNTSTGSSPIFDVNFQFDPTAEIDTGFPTFFHNSI

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>AtWRKY42 MFRFPVSLGGGPRENLKPSDEQHQRAVVDEVDFFRSAEKRDRVSREEQNIIADETHRVHVKRENSRVDDH DDRSTDHINIGLNLLTANTGSDESMVDDGLSVDMEEKRTKCENAQLREELKKASEDNQRLKEMLSQTTNN FNSLQMQLVAVMRQQEDHHHLATTENNDNVKNRHEVPEMVPRQFIDLGPHSDEVSSEERTTVRSGSPPSL LEKSSSRQNGKRVLVREESPETESNGWRNPNKVPKHHASSSICGGNGSENASSKVIEQAAAEATMRKARV SVRARSEAPMLSDGCQWRKYGQKMAKGNPCPRAYYRCTMAVGCPVRKQVQRCAEDRTILITTYEGNHNHP LPPAAMNMASTTTAAASMLLSGSTMSNQDGLMNPTNLLARTILPCSSSMATISASAPFPTITLDLTESPN GNNPTNNPLMQFSQRSGLVELNQSVLPHMMGQALYYNQQSKFSGLHMPSQPLNAGESVSAATAAIASNPN FAAALAAAITSIINGSNNQQNGNNNNSNVTTSNVDNRQ

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>AtWRKY43 MNGLVDSSRDKKMKNPRFSFRTKSDADILDDGYRWRKYGQKSVKNSLYPRCTQHMCNVKKQVQRLSKETS IVETTYEGIHNHPCEELMQTLTPLLHQLQFLSKFT

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na

>AtWRKY44 MEVNDGERVVIAKPVASRPSSSSGFRTFTELLTDSVTVSPQTTCHEIVDAAIRPKTLRFNQPVAASVSCP RAEVKGIGNGMSCDDDSDSRNYVVYKPKAKLVSKATVSALANMLQGNRQQTWRQSEAVSYGKSVSQGTHR AGPNLVQKVPSFTESETSTGDRSSVDGYNWRKYGQKQVKGSECPRSYYKCTHPKCPVKKKVERSVEGQVS EIVYQGEHNHSKPSCPLPRRASSSISSGFQKPPKSIASEGSMGQDPNNNLYSPLWNNQSNDSTQNRTEKM SEGCVITPFEFAVPRSTNSNPGTSDSGCKSSQCDEGELDDPSRSKRRKNEKQSSEAGVSQGSVESDSLED GFRWRKYGQKVVGGNAYPRSYYRCTSANCRARKHVERASDDPRAFITTYEGKHNHHLLLSPPSSSTLPFN SPQLSKQTIPPFCLKE SLDNIIDFGQDWLWS >AtWRKY45 MEDRRCDVLFPCSSSVDPRLTEFHGVDNSAQPTTSSEEKPRSKKKKKEREARYAFQTRSQVDILDDGYRW RKYGQKAVKNNPFPRSYYKCTEEGCRVKKQVQRQWGDEGVVVTTYQGVHTHAVDKPSDNFHHILTQMHIF PPFCLKE >AtWRKY46 MMMEEKLVINELELGKELANRLMNNLKHTSSVDSNKTLISDILRIYQNAIFMLSFNQDKNILKRSLEIDG KDSKNVFKKRKVSEKNTEKVKVFVATEQENGSIDDGHCWRKYGQKEIHGSKNPRAYYRCTHRFTQDCLAV KQVQKSDTDPSLFEVKYLGNHTCNNITSPKTTTNFSVSLTNTNIFEGNRVHVTEQSEDMKPTKSEEVMIS LEDLENKKNIFRTFSFSNHEIENGVWKSNLFLGNFVEDLSPATSGSAITSEVLSAPAAVENSETADSYFS SLDNIIDFGQDWLWS >AtWRKY47 MEEHIQDRREIAFLHSGEFLHGDSDSKDHQPNESPVERHHESSIKEVDFFAAKSQPFDLGHVRTTTIVGS SGFNDGLGLVNSCHGTSSNDGDDKTKTQISRLKLELERLHEENHKLKHLLDEVSESYNDLQRRVLLARQT QVEGLHHKQHEDVPQAGSSQALENRRPKDMNHETPATTLKRRSPDDVYGRDMHRGSPKTPRIDQNKSTNH EEQQNPHDQLPYRKARVSVRARSDATTVNDGCQWRKYGQKMAKGNPCPRAYYRCTMAVGCPVRKQVQRCA EDTTILTTTYEGNHNHPLPPSATAMAATTSAAAAMLLSGSSSSNLHQTLSSPSATSSSSFYHNFPYTSTI ATLSASAPFPTITLDLTNPPRPLQPPPQFLSQYGPAAFLPNANQIRSMNNNNQQLLIPNLFGPQAPPREM

Journal Pre-proof VDSVRAAIAMDPNFTAALAAAISNIIGGGNNDNNNNTDINDNKVDAKSGGSSNGDSPQLPQSCTTFSTN >AtWRKY48 MEKKKEEDHHHQQQQQQQKEIKNTETKIEQEQEQEQKQEISQASSSSNMANLVTSSDHHPLELAGNLSSI FDTSSLPFPYSYFEDHSSNNPNSFLDLLRQDHQFASSSNSSSFSFDAFPLPNNNNNTSFFTDLPLPQAES SEVVNTTPTSPNSTSVSSSSNEAANDNNSGKEVTVKDQEEGDQQQEQKGTKPQLKAKKKNQKKAREARFA FLTKSDIDNLDDGYRWRKYGQKAVKNSPYPRSYYRCTTVGCGVKKRVERSSDDPSIVMTTYEGQHTHPFP MTPRGHIGMLTSPILDHGATTASSSSFSIPQPRYLLTQHHQPYNMYNNNSLSMINRRSSDGTFVNPGPSS SFPGFGYDMSQASTSTSSSIRDHGLLQDILPSQIRSDTINTQTNEENKK >AtWRKY49 MEEEGYQWTRRCGNNAVEDPFVYEPPLFFLPQDQHHMHGLMPNEDFIANKFVTSTLYSGPRIQDIANALA LVEPLTHPVREISKSTVPLLERSTLSKVDRYTLKVKNNSNGMCDDGYKWRKYGQKSIKNSPNPRSYYKCT NPICNAKKQVERSIDESNTYIVTYEGFHFHYTYPFFLPDKTRQWPNKKTKIHKHNAQDMNKKSQTQEESK EAQLGELPNQNHPVNKAQENTPANLEEGLFFPVDQCRPQQGLLEDVVAPAMKNIPTRDSVLTAS

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>AtWRKY50 MNDADTNLGSSFSDDTHSVFEFPELDLSDEWMDDDLVSAVSGMNQSYGYQTSDVAGALFSGSSSCFSHPE SPSTKTYVAATATASAGKFLFWFLTIRTDHRKFKNISP

-p

ro

>AtWRKY51 MNISQNPSPNFTYFSDENFINPFMDNNDFSNLMFFDIDEGGTNGLIEEEISSPTSIVSSETFTGESGGSG SATTLSKKESTNRGSKESDQTKETGHRVAFRTRSKIDVMDDGFKWRKYGKKSVKNNINKRNYYKCSSEGC SVKKRVERDGDDAAYVITTYEGVHNHESLSHVYYNEMVLSYDHDNWNQHSLLRS

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na

lP

re

>AtWRKY53 MEGRDMLSWEQKTLLSELINGFDAAKKLQARLREAPSPSSSFSSPATAVAETNEILVKQIVSSYERSLLL LNWSSSPSVQLIPTPVTVVPVANPGSVPESPASINGSPRSEEFADGGGSSESHHRQDYIFNSKKRKMLPK WSEKVRISPERGLEGPQDDVFSWRKYGQKDILGAKFPRSYYRCTHRSTQNCWATKQVQRSDGDATVFEVT YRGTHTCSQAITRTPPLASPEKRQDTRVKPAITQKPKDILESLKSNLTVRTDGLDDGKDVFSFPDTPPFY NYGTINGEFGHVESSPIFDVVDWFNPTVEIDTTFPAFLHESIYY >AtWRKY54 MDSNSNNTKSIKRKVVDQLVEGYEFATQLQLLLSHQHSNQYHIDETRLVSGSGSVSGGPDPVDELMSKIL GSFHKTISVLDSFDPVAVSVPIAVEGSWNASCGDDSATPVSCNGGDSGESKKKRLGVGKGKRGCYTRKTR SHTRIVEAKSSEDRYAWRKYGQKEILNTTFPRSYFRCTHKPTQGCKATKQVQKQDQDSEMFQITYIGYHT CTANDQTHAKTEPFDQEIIMDSEKTLAASTAQNHVNAMVQEQENNTSSVTAIDAGMVKEEQNNNGDQSKD YYEGSSTGEDLSLVWQETMMFDDHQNHYYCGETSTTSHQFGFIDNDDQFSSFFDSYCADYERTSAM >AtWRKY55 MEEVMSMIFHGMKLVKSLESSLPEKPPESLLTSLDEIVKTFSDANERLKMLLEIKNSETALNKTKPVIVS VANQMLMQMEPGLMQEYWLRYGGSTSSQGTEAMFQTQLMAVDGGGERNLTAAVERSGASGSSTPRQRRRK DEGEEQTVLVAALRTGNTDLPPDDNHTWRKYGQKEILGSRFPRAYYRCTHQKLYNCPAKKQVQRLNDDPF TFRVTYRGSHTCYNSTAPTASSATPSTIPISSVTTGHSVDYGLAVVDMADVMFGSGGVGTNMDFIFPKND PS >AtWRKY56 MEGVDNTNPMLTLEEGENNNPFSSLDDKTLMMMAPSLIFSGDVGPSSSSCTPAGYHLSAQLENFRGGGGE MGGLVSNNSNNSDHNKNCNKGKGKRTSAMQRIAFHTRSDDDVLDDGYRWRKYGQKSVKNNAHPRSYYRCT YHTCNVKKQVQRLAKDPNVVVTTYEGVHNHPCEKLMETLSPLLRQLQFLSRVSDL >AtWRKY57 MNDPDNPDLSNDDSAWRELTLTAQDSDFFDRDTSNILSDFGWNLHHSSDHPHSLRFDSDLTQTTGVKPTT VTSSCSSSAAVSVAVTSTNNNPSATSSSSEDPAENSTASAEKTPPPETPVKEKKKAQKRIRQPRFAFMTK SDVDNLEDGYRWRKYGQKAVKNSPFPRSYYRCTNSRCTVKKRVERSSDDPSIVITTYEGQHCHQTIGFPR GGILTAHDPHSFTPHHHLPPPLPNPYYYQELLHQLHRDNNAPTPRLPRPTTEDTPVVSNPTEEGLLGDIV PQTMRNP >AtWRKY58 MAVEDDVSLIRTTTLVAPTRPTITVPHRPPAIETAAYFFGGGDGLSLSPGPLSFVSSLFVDNFPDVLTPD NQRTTSFTQLLNGTMSVSPGGGGRSTAGMFAGGGPMFTIPSGFSPSSLLTSPMFFPPQSSAHTGFIQPRQ

Journal Pre-proof QSQPQPQRPDTFPHHMPPSTSVAVHGRQSLDVSQVDQRARNHYNNPGNNNNNRSYNVVNVDKPADDGYNW RKYGQKPIKGCEYPRSYYKCTHVNCPVKKKVERSSDGQITQIIYKGQHDHERPQNRRGGGGRDSTEVGGA GQMMESSDDSGYRKDHDDDDDDDDEDDEDLPASKIRRIDGVSTTHRTVTEPKIIVQTKSEVDLLDDGYRW RKYGQKVVKGNPHPRSYYKCTTPNCTVRKHVERASTDAKAVITTYEGKHNHDVPAARNGTAAATAAAVGP SDHHRMRSMSGNNMQQHMSFGNNNNTGQSPVVLRLKEEKITI >AtWRKY59 MNYPSNPNPSSTDFTEFFKFDDFDDTFEKIMEEIGREDHSSSPTLSWSSSEKLVAGEITSPLQTSLATSP MSFEIGDKDEIKKRKRHKEDPIIHVFKTKSIDEKVALDDGYKWRKYGKKPITGSPFPRHYHKCSSPDCNV KKKIERDTNNPDYILTTYEGQHNHPSPSVVYCDSDDFDLNSLINNWSFQTANTYSFSHSAPY

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>AtWRKY60 MDYDPNTNPFDLHFSGKLPKREVSASASKVVEKKWLVKDEKRNMLQDEINRVNSENKKLTEMLARVCEKY YALNNLMEELQSRKSPESVNFQNKQLTGKRKQELDEFVSSPIGLSLGPIENITNDKATVSTAYFAAEKSD TSLTVKDGYQWRKYGQKITRDNPSPRAYFRCSFSPSCLVKKKVQRSAEDPSFLVATYEGTHNHTGPHASV SRTVKLDLVQGGLEPVEEKKEKGTIQEVLVQQMASSLTKDPKFTAALATAISGRLIEHSRT

re

-p

ro

>AtWRKY61 MEKDDFLRSGHGREESHDEMRKLDSSHDDSHQEHDHIIRSKLDSTKVEMDEAKEENRRLKSSLSKIKKDF DILQTQYNQLMAKHNEPNKFQSKGHHQDKGEDEDREKVNEREELVSLSLGRRLNSEVPSGSNKEEKNKDV EEAEGDRNYDDNEKSSIQGLSMGIEYKALSNPNEKLEIDHNQETMSLEISNNNKIRSQNSFGFKNDGDDH EDEDEILPQNLVKKTRVSVRSRCETPTMNDGCQWRKYGQKIAKGNPCPRAYYRCTIAASCPVRKQVQRCS EDMSILISTYEGTHNHPLPMSATAMASATSAAASMLLSGASSSSSAAADLHGLNFSLSGNNITPKPKTHF LQSPSSSGHPTVTLDLTTSSSSQQPFLSMLNRFSSPPSNVSRSNSYPSTNLNFSNNTNTLMNWGGGGNPS DQYRAAYGNINTHQQSPYHKIIQTRTVGSSFDPFGRSSSSHSPQTNLDHIGIKNIISHQVPSLPAETIKA ITTDPSFQSALATALSSIMGGDLKIDHNVTRNEAEKSP

na

lP

>AtWRKY62 MNSCQQKAMEKLLHGHGCANQLLIMDQTESDSSMEREDLAKSVLHCFSDALSILIDTNDHQDDQSNNSSP QDSSPVLESSRKPLHKRGRKTSMAESSDYHRHESSTPIYHDGFLWRKYGQKQIKESEYQRSYYKCAYTKD QNCEAKKQVQKIQHNPPLYSTTYFGQHICQLHQAYATFPIDTSDFEEHEGSHMIRFGHPNISFSSSTSNL RQHQNHQDRIKDEYMKPVIAEDWSPSQWMSSEVALAVEAFEFNPFWTSHDLSS

Jo ur

>AtWRKY63 MFSNIDHKAVAALLHGQGCANILKTVLDNCKVSSVSTEPLINTILDSFSLALSSVNSPNRQPHHESSSRD MAGLVPQRSSKKKICGVKGLEIYRDDSPNPRLDDGFTWRKYGQKTIKTSLYQRCYYRCAYAKDQNCYATK RVQMIQDSPPVYRTTYLGQHTCKAFGVHDNTYGSEMINFDQVVSESVMRQLATIGEQAVLMEDEANHIMN QEYDINDYLVDDEVFWGNEFPLFSSEDLMLF >AtWRKY64 MFSNIDQTAVAALLRGQGCANSLKRLLENHKLSSDSTEPLIYTILNSFSLALSFVDPPSLLPHNESSLQN MTSHVLQRSSKKKYYGAEDLEYYRDESPTPRPDDGFTWRKYGQKTIKTSPYQRCYYRCTYAKDQNCNARK RVQMIQDNPPVYRTTYLGKHVCKAVAVHDDTYGSEMIKFDQVVSESVMPQLATIDEQAITMEDEAIDHIM NQECDINDFSVDDDPFWASQFPPFSSEDIMFFDNIANLD >AtWRKY65 MKRGLDMARSYNDHESSQETGPESPNSSTFNGMKALISSHSPKRSRRSVEKRVVNVPMKEMEGSRHKGDT TPPSDSWAWRKYGQKPIKGSPYPRGYYRCSSTKGCPARKQVERSRDDPTMILITYTSEHNHPWPLTSSTR NGPKPKPEPKPEPEPEVEPEAEEEDNKFMVLGRGIETTPSCVDEFAWFTEMETTSSTILESPIFSSEKKT AVSGADDVAVFFPMGEEDESLFADLGELPECSVVFRHRSSVVGSQVEIF >AtWRKY66 MSLEIDAKAVSALLLGQGCANNLKTLLKNHETGSVSTEPLINSILDSFSFALSSQNIPRHVSQRSSKNIL VNIALIQHCRKMCGIQGIEDSPTPAHIDGFIWRKYGQKTIKTSPHQRWYYRCAYAKDQNCDATKRVQKIQ DNPPVYRNTYVGQHACEAPAYAVNNGTYGSKMIKFDYVMPESVMPQPLSIDSQEITMEDKDTDDHILNYI NEHLMEDEAYDVFPDVLGERCCFGLEPFPGLNIYKS >AtWRKY67 MVSNIDHKAMEALLRGQGCANSLKILLENGEISSVSTKPLINTILDSFSFALSFVDSPNHPPYHESSSQN

Journal Pre-proof MASHMSRRSSKQKLCVAEGLVNYRDDSPIMCPNDGFTWRKYGQKTIKGSLHKRCYYRCTYAKDQNCNATK QVQKIKDNPPVYRTTYLGKHVCKAVAVHDDTYGSTMIRFDQDVSESIMPQLTTIDHQAITMEENSADHIM NQECDINDYLVGYDPFWASQFPPFPSSDTMFLENIAAFD >AtWRKY68 MENIGVGMPFYDLGQTRVYPLLSDFHDLSAERYPVGFMDLLGVHRHTPTHTPLMHFPTTPNSSSSEAVNG DDEEEEDGEEQQHKTKKRFKFTKMSRKQTKKKVPKVSFITRSEVLHLDDGYKWRKYGQKPVKDSPFPRNY YRCTTTWCDVKKRVERSFSDPSSVITTYEGQHTHPRPLLIMPKEGSSPSNGSASRAHIGLPTLPPQLLDY NNQQQQAPSSFGTEYINRQEKGINHDDDDDHVVKKSRTRDLLDGAGLLKDHGLLQDVVPSHIIKEEY >AtWRKY69 MHRRAAIQESDDEEDETYNDVVPESPSSCEDSKISKPTPKKSRRNVEKRVVSVPIADVEGSKSRGEVYPP SDSWAWRKYGQKPIKGSPYPRGYYRCSSSKGCPARKQVERSRVDPSKLMITYACDHNHPFPSSSANTKSH HRSSVVLKTAKKEEEYEEEEEELTVTAAEEPPAGLDLSHVDSPLLLGGCYSEIGEFGWFYDASISSSSGS SNFLDVTLERGFSVGQEEDESLFGDLGDLPDCASVFRRGTVATEEQHRRCDFGAIPFCDSSR

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>AtWRKY70 MDTNKAKKLKVMNQLVEGHDLTTQLQQLLSQPGSGLEDLVAKILVCFNNTISVLDTFEPISSSSSLAAVE GSQNASCDNDGKFEDSGDSRKRLGPVKGKRGCYKRKKRSETCTIESTILEDAFSWRKYGQKEILNAKFPR SYFRCTHKYTQGCKATKQVQKVDLEPKMFSITYIGNHTCNTNAETPKSKTCDHHDEIFMDSEDHKSPSLT TSMKEEEDNPHRHHGSSTENDLSLVWPEMVFEEDYHHQASYVNGKTSTSIDVLGSQDLMVFGGGGDFEFS ENEHFSIFSSCSNLS

lP

re

-p

>AtWRKY71 MDDHVEHNYNTSLEEVHFKSLSDCLQSSLVMDYNSLEKVFKFSPYSSPFQSVSPSVNNPYLNLTSNSPVV SSSSNEGEPKENTNDKSDQMEDNEGDLHGVGESSKQLTKQGKKKGEKKEREVRVAFMTKSEIDHLEDGYR WRKYGQKAVKNSPYPRSYYRCTTQKCNVKKRVERSFQDPSIVITTYEGKHNHPIPSTLRGTVAAEHLLVH RGGGGSLLHSFPRHHQDFLMMKHSPANYQSVGSLSYEHGHGTSSYNFNNNQPVVDYGLLQDIVPSMFSKN ES

Jo ur

na

>AtWRKY72 MEVLLKLPSSESPLKDKFGSVQIHEANKGDGDHQELESAKAEMSEVKEENEKLKGMLERIESDYKSLKLR FFDIIQQEPSNTATKNQNMVDHPKPTTTDLSSFDQERELVSLSLGRRSSSPSDSVPKREEKTDAISAEVN ADEELTKAGLTLGINNGNGGEPKEGLSMENRANSGSEEAWAPGKVTGKRSSPAPASGGDADGEAGQQNHV KRARVCVRARCDTPTMNDGCQWRKYGQKIAKGNPCPRAYYRCTVAPGCPVRKQVQRCADDMSILITTYEG THSHSLPLSATTMASTTSAAASMLLSGSSSSPAAEMIGNNLYDNSRFNNNNKSFYSPTLHSPLHPTVTLD LTAPQHSSSSSSSLPSLNFNKFSNSFQRFPSTSLNFSSTSSTSSNPSTLNLPAIWGNGYSSYTPYPYNNV QFGTSNLGKTVQNSQSLTETLTKALTSDPSFHSVIAAAISTMVGSNGEQQIVGPRHSISNNIQQTNTTNN NKGCGGYFSSLLMSNIMASNQTGASLDQPSSQLPPFSMFKNSSSSSSTTNFVNKEEKS >AtWRKY74 MEEVEAANKAAVESCHGVLNLLSQQTNDSKSIMVETREAVCKFKRVSSLLSRGLGQRKIKKLNNNNYKFS SSLLPQHMFFESPVCSNNAISGCIPILAPKPLQIVPAGPPPLMLFNQKMCLDKSFLELKPPSSRAVDPKP YQFIHTHQQGVYSRSKSGLNLKFDGSIGASCYSPSISNGSRSFVSSLSMDGSVTDYDRKSFHLIGLPQGS DHISQHSRRTSCSGSLKCGSKSKCHCSKKRKLRVKRTIKVPAISNKIADIPPDEYSWRKYGQKPIKGSPH PRGYYKCSSVRGCPARKHVERCVEETSMLIVTYEGEHNHSRILSSHSAHT >AtWRKY75 MEGYDNGSLYAPFLSLKSHSKPELHQGEEESSKVRSEGCSKSVESSKKKGKKQRYAFQTRSQVDILDDGY RWRKYGQKAVKNNKFPRSYYRCTYGGCNVKKQVQRLTVDQEVVVTTYEGVHSHPIEKSTENFEHILTQMQ IYSSF

>OsWRKY1 MDKGHLGGGGGGGGGLLALDASPRPLGFLNLLSPPPFHRSTMEADDSGGGGGRARRSVEVDFFSDEKKNM KKSRVSGGVAAEADDAKGPAAAGLAIKKEDLTINLLPAGNNARSDRSMVVDDDAASRPDHEEKSRSSNEL AAMQAELGRMNEENQRLRGMLTQVTTSYQALQMHLVALMQQRPQMMQPPTQPEPPPPHQDGKAEGAVVPR QFLDLGPSSGAGGEAAEEPSNSSTEAGSPRRSSSTGNKDQERGDSPDAPSTAAAWLPGRAMAPQMGAAGA AGKSHDQQAQDANMRKARVSVRARSEAPIIADGCQWRKYGQKMAKGNPCPRAYYRCTMATGCPVRKQVQR

Journal Pre-proof CAEDRSILITTYEGTHNHPLPPAAMAMASTTSAAASMLLSGSMPSADGAAGLMSSNFLARTVLPCSSSMA TISASAPFPTVTLDLTHAPPGAPNAVPLNAARPGAPAPQFQVPLPGGGMAPAFAVPPQVLYNQSKFSGLQ MSSDSAEAAAAAAAAAQFAQPRPPIGQLPGPLSDTVSAAAAAITADPNFTVALAAAITSIIGGQHAAAAG NSNANNTNTNTTSNTNNTSSNNTTSNNTNSETQ

>OsWRKY2 MCDSLFWQSSADQGDLSDVVRASLQLQTAPRHQAASPPYVHLLGGGGGGGEDQLAAVSQHAEQQQQSMVD ASAACDLLHALLPPPPVVQVQQQGASRTRTTIEEDTTGDGEELFAGAHYVVPPIKRRKSQTKKVVCIPAG ASGGGGGEVVPSDLWAWRKYGQKPIKGSPYPRGYYRCSSSKGCSARKQVERSRADPTMLVVTYTSDHNHP WPTHRNALAGSTRPSSSNSSNIRLQDSTPVHHQSQTGHDRLTTTHLKQEDVIISPSLLQPDHHQLCTIID TKHHLLFHQDYPHSFGLFD

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>OsWRKY3 MAGAEWSPFDGDAAFAEYSSAVLAELGGWAAPGEEGGAGMMVPAALDLPVDVVGAAAREEEEEEEEAPAR SGDGAAAAASSSSSGEPAAPDKRPAAAEAAPAAAATATATAKKGQKRARQPRFAFMTKSEIDHLEDGYRW RKYGQKAVKNSPFPRSYYRCTNSKCTVKKRVERSSDDPSVVITTYEGQHCHHTASFQRGVGGAAVAAHIH GAAAVALAEQMSAFVSPPPQPHMLYGLPRLHPPSSETAVSCSMPTTTSLQELNNSEGLQRPGYNNSPQAA VTIAQRPPSPSVPPAVSFDKGLLDDIVPPGVRLG

lP

re

-p

>OsWRKY4 MADRRRSDGGGGMQQQPFTSPGQERVFDGGGVPGQVAAPYGSDFDQSSYMALLAAGAVGVGVGVQPTAAP WAVEEDVAAAPPGISLAPQFSMANYAPPPSYQHPATLVSPPLAAGLHPYPPYLHGVDAPPPQWPPRPAPP PSFSVLDLAAAAAPHEQRHSMQQLLLRAAAFGGGMHAAAAPAPAAAAAIEQPAKDGYNWRKYGQKQLKDA ESPRSYYKCTRDGCPVKKIVERSSDGCIKEITYKGRHSHPRPVEPRRGGAASSSSSAMAAGTDHNAGAAA DDAAAADEDDPSDDDDTLLHEDDDDGEEGHDRGVDGEVGQRVVRKPKIILQTRSEVDLLDDGYRWRKYGQ KVVKGNPRPRSYYKCTADGCNVRKQIERASADPKCVLTTYTGRHNHDPPGRPPAAANLQMPGPAAMRLAG GGTAHQQPSGGAHQMKEET

Jo ur

na

>OsWRKY5 MEMMVQKQRHEEGEEERGGLCAREIKELDFFSAAGAGAGRRDDDDVLRADGISSSHAGFMVSTALDLLTA VNDGDHHEEKKGQSNIHQSKQMDAAATTVEGELRQAGEENRRLRRRLEELTSSYGALYHQLVQAQQLHTK HQQQAPIAGVQLLDALAAASPASHRRRAAAAVDGDRTADSDGGEGDENVSPSLGSKRPAAAATLTRLTPE SGSGGENNGGGEQAPAAEMAPCRKARVSVRARSEAPMISDGCQWRKYGQKMAKGNPCPRAYYRCTMASQC PVRKQVQRCAKDKSILITTYEGTHSHPLPPAAAAMAKTTSAAAAMLLSGPAVSRDALFAAHHHVVAPPPF FHHPYAGSTMATLSASAPFPTITLDLTQPPPTTTTTAAAAMLQLHRPYAFSSLPFSMYGAGGGSHRPPVV LPPPSSVVETMTAAITRDPNFTTAVAAALSSIMAGGGAQARTPPRGGSDAAGDINGGGGADHATAGARAA AAATQPCGTSPT

>OsWRKY6 MDGVEEANMAAVESSKKLVAILSKSGDPFRLMAAVAETDEAVSRFGKVVTILSNRVGHARARLGKRRSSP PVDPGCLMDHPLAAAASSPAPSNGRLHFSSSAATASPSPATAAAASSAANVTPAVVDRSLFLETTLLDLN SRGAPAPAASMAAAAKNSSKLAPAPMVNSSSSANHIQFQQPMKSFQFEQTPISDKFHIEMPRGVGGGGGK EVISFSFDNSVCTSSAATSFFTSISSQLISMSDAATNSAAAAAAPTTKKPSSCARKATADDDAGGKCHCP KKKKPREKKVVTVPAISDKVADIPSDNYSWRKYGQKPIKGSPHPRGYYRCSSKKDCPARKHVERCRSDPA MLLVTYENEHNHAQPLDLSVVQQATANPQT

>OsWRKY7 MAAVGAHAAVYHHPVSGLSAPAGDAAYSMSSYFSHGGSSTSSSASSFSAALAAATTPPLPDPSGSQFDIS EFFFDDAPPAAVFNGAPTAALPDGAAANATRSAAEAVPAPAPAAVERPRTERIAFRTKSEIEILDDGYKW RKYGKKSVKNSPNPRNYYRCSTEGCNVKKRVERDKDDPSYVVTTYEGTHNHVSPSTVYYASQDAASGRFF VAGTQPPGSLN

Journal Pre-proof >OsWRKY8 MSGPGGGGHGLYEDHPAAAGFLPFDHDDDVVASFFFGRSAASGGGAGAGAGAGDDDGVGLITPYSSITDY LQGFLQDPVYASSPLGGDAAVKHETVVDHPSQAGGVAAAPATPNSSVLSSSSEAAGGDDLRRCKKGRRPE DEEEEEIDDEGSAVQSCKTNKMKNKKGAKKEREPRVAFMTKSEVDHLEDGYRWRKYGQKAVKNSSYPRSY YRCTAPRCGVKKRVERSEQDPSMVITTYEGQHTHPSPVSYHMHRQQGLMHVSARGVMPGAAGAYQFGAPP PPLLGFDEALAARVRMTMNQQQQQQQLGFVPSIHAAAARPTMPPLHLYTAQQDLFLP

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>OsWRKY9 MELRPPPPKHHHHRRRRGGGGEDGGEEEEEETGRLSLRGGGFWRRHDGEEEEEKGGGRRGEIKEVDFFLG ASGRDVVVASRRHDDGFRGTTHGGGGGGDVNIGLDLLTTTTAGAAAGGAAAGAGEEDTGKNHRKEATTAA VDVELRRVVEENRRLRGMLDELNRSYSALYHQYLQVTQQQNHRHPDHHLIMNNNNNRPSLAQTHRTAATT TATTQQFLEPRASSTAQATADADMAASDDEAGRGGGDGDASSPSLSNAAGGGGGGNKMRRVGGQDETAAA APARENGEQQAAAAAELPCRKPRVSISDGCQWRKYGQKMAKGNPCPRAYYRCTMAIGCPVRKQVQRCAED KTVLITTYEGNHNHQLPPAATTMANTTSAAAAMLLSGPAASRDGAAAALLGHHHHHHPAAMFHQSFPYAS TMATLSASAPFPTITLDLTQTPAGGAGAASLLHALHRPPVIHPGAAAQAMPFAVPPQLAMYLPQQRAAAA GLGGAGAARQPSVMETVTAALAADPNFTTALAAAISSVVAGGAHHQALSTTPRGSAAGAGDGNGNGSSAA AVATGAASPAATAEAPAASGSPPRLATQSCTTSN

re

-p

>OsWRKY10 MAASLGLCHETSYAYSYPASNTSSSLCFPPLMADHIVDGGGGGGCSFGEFLELGHSVYSLPLPPPPSQPV VVAGGNNDQYGVSSSSSAAATTSRIGFRTRSEVEVLDDGFKWRKYGKKAVKSSPNPRNYYRCSAAGCGVK KRVERDGDDPRYVVTTYDGVHNHATPGCVGGGGHLPYPTSAAPPWSVPAAAASPPPAHAQAWGAPLHAAA AAHSSESPMQEHFD

na

lP

>OsWRKY11 MSSGGGGGGGGDRHGPYHQHGHLGRGEGADYVYSSSDMESFFFSQPGGVGIGGGGGGVVGAGGADEIMPY SSITDYLQGLLDPSGLARHLDVACPSSQDTVVKQELSVDVTSHDSQGTGGVAGEGVAQATPNSSASFSSS DGEAEGGKSSRRCKKGQAKAEEEDDKDEEDGENSKKPNKPKKKAEKRQRQPRVAFLTKSEVDHLEDGYRW RKYGQKAVKNSPYPRSYYRCTTPKCGVKKRVERSYQDPSTVITTYEGQHTHHSPASLRGGGGGVGIVGGH HHHHLFMPGVHGLPPSHLMPAGFHPELMGLMHHHPAMAAAAANPSMYFPGVAASAPPPPAVAGGGAMPPN DHPPLQQHHFTDYALLQDLFPSTMPSSNP

Jo ur

>OsWRKY12 MRGPLLLRAVVVVAMEHFNDWDLQAVVRSCSFPQSEPPRVGVGVPAAPGAGGAPVVVAPPARAPDGPDQM ARASASALYDLEYLDLDHKPFLLPGSSSSSSSSRAVARARGEDDGKGRHEVMISFPAAAAASTSGAQPRS PSGRKPGIRTPRPKRRKFSAYFEVIDYCDFHGVRSSKKSQLKKVVYEVPVADGGVSSDLWAWRKYGQKPI KGSPYPRGYYKCSSMKGCMARKMVERSPAKPGMLVVTYMAEHCHPVPTQLNALAGTTRHKSAPTGDDDKP TSPGPAAGRAAAGEGVVKCEDVDGNELSAMAADGGAEDTAAAADDDGELWPEGMGLELDEFLGPMDDDVF EFDHVLEDDGVLGRRLSL

>OsWRKY13 MAAGEEVMDRSTSAEDGYCSAGTDSPRAESVDEQGAAEESSPRGGQKRELPSPSASPSSPLPPAAKRSRR SVEKRVVSVPIAECGDRPKGAGEGPPPSDSWAWRKYGQKPIKGSPYPRGYYRCSSSKGCPARKQVERSRA DPTVLLVTYSFEHNHPWPQPKSSSCHASKSSPRSTAPKPEPVADGQHPEPAENESSASAELEVPEPEPEQ ESEPVVKQEEEQKEEQKAVVEPAAVTTTVAPAPAVEEEDENFDFGWIDQYHPTWHRSYAPLLPPEEWERE LQGDDALFAGLGELPECAVVFGRRRELGLAATAPCS

>OsWRKY14 MDGEWSDGAAVSSPTMSGGGGREQMKGGEDVAAADCPGSPVSPSPAAAQRSAAGAAASPSGRSRRSAQKR VVTVPLADVTGPRPKGVGEGNTPTDSWAWRKYGQKPIKGSPFPRAYYRCSSSKGCPARKQVERSRNDPDT VIVTYSFEHNHSATVPRAQNRQAAPQKPKAQACSPPEPVVEVEPEETHQYGVTAGPATGGGGGAAAIEVR DEFRWLYDVVSVPATSTSPSDIDAADEMQLYDQPMFFGGAVVGTAALLPDEFGDVGGLGGEGLGEEEALF EGLGELPECAMVFRRRAGDGLEMGGGVKIEQPAESTAMT

Journal Pre-proof

>OsWRKY15 MEGSSSDGSRYGGSGRCALVAELVQMQGMVRQLEAEMGTHGGGGGGGGGGGADGRRSGRAGAARCIARSC CTEAAAAGRLTQQAGAAPESPPSADGSAGSDLGADSRCRANAAGPCKKRKTLPKWSKQVKVRSVQDVGPL DDGFSWRKYGQKDILGAKYPRAYFRCTHRHTQGCHASKQVQRADGDPLLFDVVYHGDHTCAHGVRSAAAA IDGQAAASAEQKHQPTPPQEQNAVSVAFTSMAVVNASTSSPFVSPAMSDCQISYELGGGSMAGVRNVPDV ELASKTNSSMGDDMEFMFSLDSDFLDTYKYSSYF

ro

of

>OsWRKY16 MGGEPRARAEAAWAAALPAALVALVRDTATYTRMMHRLRMETPRGPTCQLDPCCCCCTAHVIPPPPPVSR THATRRDAEASVPPPPASAAVSSRSDGTGQMAAGVTLACAAPPPLRAPRASDGGRRRGVVKGGAGTDTCR SPQRLNVRPRERERVRACVRARAKNHEHGQRRREAAVDPAMSGEYQFQDELAPLFARPGGGAGEMQMLPS SWFADYLQAGTPMQMDYDLMCRALELPVGEDVKREVGVVDVVAAGGGGAPPLTPNTTSSMSTSSSEGVGG GGGGGAGAGAGEEESPARCKKEEDENKEEGKGEEDEGHKNKKGSAAKGGKAGKGEKRARQPRFAFMTKSE VDHLEDGYRWRKYGQKAVKNSPYPRSYYRCTTQKCPVKKRVERSYQDPAVVITTYEGKHTHPIPATLRGS THLLAAHAQAAAAAAAAHQLHHHHGHHGHHGMAPPLPLGSGAAAQFGRSSGIDVLSSFLPRAAAAHHGMT TMGGAAATTTTSHGLNSAISGGGGVSSETTSAVTVAASAQPSSPAALQMQHFMAQDLGLLQDMLLPSFIH GTNQP

lP

re

-p

>OsWRKY17 MQESREYWRDGGDVVGEELLREILDETAAVHSNSNSNSNSNSNSKEAEEEDEREYFAAAAADEQLQVEAP CGRRRRESMVNKLISTVYSGPTISDIESALSFTAAGDHQLLADGHNFAASSCSPVVFSPEKTLSKTMENK YTLKMKSCGNNGGLADDGYKWRKYGQKSIKNSPNPRSYYRCTNPRCNAKKQVERAVDEPDTLIVTYEGLH LHYTYSHFLHSTSSSSSSTTTQQQLQPQPQMMTNCKKKPKLHLHPLLHDDPPPPPPPPEMTTMMIMQSFS IQQQQHDDDQLLQPAADDHLMVQAPPDDCYNINGSSSSGLMMSLDDDEQAAGAGGLLEDVVPLLVRRPPP PICNNNNYYYSPATTCTSDNEYGSSASASPSSSVSVSSWTTPMSPCIDMAILSNIF

Jo ur

na

>OsWRKY18 MASRHQLQTMQFTDPASRSPRPVGGGVHGQPPPTPMSSPFSSRKPRMQEGHPTCVNLTPIPHTDGHLWRK YGEKKIKNSSFPRLYYRCSYRDDRNCMATKVVQQENDADPPLYRVTYIHPHTCNPSPPAPTPAHVFTEPP PAKAEVHHAVLFRFSSTAGGHTANNAVHRQQWQPAAATMAAGAQAQLSMTMSDDEREQPPAAIRSAPPAR RLSMFRAVVDGLRQMRSSAPPTPSSSMVVDDGWDTFSSFDLDTCEFSVDDELLCGDHMYFPDSMQQ

>OsWRKY19 MVELCGGEGEGQIMLATELAQLRAMARELEAKMDPDRVAARELCRALASSVDRSIRLAASCFPPPEHPPP AAGNAGRDAAFKKRQVMVPGHGGLISGKGMAKVRRQVRVTSVQDTASLDDGLSWRKYGQKDILGAKYPRA YFRCTHRHTQGCNATKQVQRADGDPLLFDVVYLGDHTCGQAAVAAAAQSAPPEHAGQEQQRQSSLLAAGT EGIHQQVVAEPMAAPFLFTSTAAGGVDDGYFSFISPANSDCQFSSDFSAGSVGVDMDHEARFEDLFSSTL EFFQSEIQNL >OsWRKY20 MAFGQDTIEQLYRELAGGRRLSAKLQALLEGPLDSRGQKEAVDVSRELGRVFMVSLYMLKPCSNSSRRPE GVTRTAPETRTDDSICLHTPARVKRVRSEEVLVRNGREEVVTRTEIITPSPYKDGYQWRKYGQKNIQDSN YLRLYFKCTFSRERSCAAKKQVQQRDAGEPPMFLVTYLNEHTCQQPQAVPGTPNTAGSSPTTTSRQRQSS SSPPAEMLDLTMNGAGLFSRLLLPHAVGGGGSAAEEEAAIVTCLAAVISGGGAAAAPPPLIWPTSAPEAA FVASAAGHSPSAADESVADEAAAAQMADMDYCFGQYDQSTFGAAAAADHRVLIGDDGDVQRIVAARIADT VWPRYTRDTSAWETAGTSSMRGSID >OsWRKY21 MAMLGSSSAVVLELMTMGYQSAAYLGELLRAASPAQAGDEQQELAAEILRCCDRVIAKLNRGGATGATTG KKRKAAESAAAAAVTSPSLPVTPTKRRARGAEAVREVRSGTTTDGFIWRKYGQKEINGCKHPRLYYRCAF RGQGCLATRRVQQSQSQDDPAAAFVIAYYGEHTCGGDAAAAAACRDGELMPPAVINSGASSFAAAWNMAS

Journal Pre-proof REPASSLAVERRSCDGDAPSETSQGWSPSFSSEVELDVVGFDLAGADSSASPVWEFLNGSFDWEFVINSL >OsWRKY22 MVKRSDNMDSSSECSRGAHKRLLQDSRSYDQENAMKKVCIGTRTEYTYAPYHDGYQWRKYGQKMIRGNSF PRCYYRCTYHQDHGCPASKHVEQHNSEDPPLFRVIYTNEHTCGTSNSASDYMASSMQIQQIADASLRKAQ AAERLRKAEVETPRLMHSPPPRCSGGYNMAMKEEKDVIVSSLLTVIRGCHIAESAGNNSAAALPVNRPPP AVARSDHYSCSYAISPELLPASDDLTLDFMLDSVLDPHWVEPLDLAWLKESTHTG >OswRKY23 MENLQLQGDDHDDEALPHFPYFAVPSPPPLAVAPAASATTSDGHQHGPLEVLEQPPCSNNLHPDGLVDGP QLAATTAVPMMLPAMTSLDWQSLLQTCLQVPPPVLEQQQPAAAAQADQYSGENDHGDLQAAESSGAGNKE KQVMAKGGAGRPSGTKKKASRPRFAFQTRSDNDILDDGYRWRKYGQKAVKNSKHPRSYYRCTHHTCNVKK QVQRLAKDTSIVVTTYEGVHNHPCEKLMEALTPILKQLQFLSQF

re

-p

ro

of

>OsWRKY24 MTTSSSGSVETSANSRLGTFSFASASFTDLLGGNAGAGGGGVSRYKAMTPPSLPLSPPPVSPSSFFNSPI GMNQADFLGSPVLLTSSIFPSPTTGAFASQHFDWRPEVAAAQSADQGGKDEQRNSYSDFSFQTAPASEEA VRTTTFQPPVPPAPLGDEAYRSQQQQQPWGYQQQPAGMDAGANAASFGAAPFQATSSEMAPQVQGGGGYS QPQSQRRSSDDGYNWRKYGQKQVKGSENPRSYYKCTFPNCPTKKKVERSLDGQITEIVYKGTHNHAKPQN TRRNSGSSAAQVLQSGGDMSEHSFGGMSGTAATPENSSASFGDDEIRVGSPRAGNGGGDEFDDDEPDSKR WRKDGDGEGISMAGNRTVREPRVVVQTMSDIDILDDGYRWRKYGQKVVKGNPNPRSYYKCTTAGCPVRKH VERASHDLRAVITTYEGKHNHDVPAARGSAALYRPAPPAAAATSSHPYLPNQPPPMSYQPTGPQPYALRP DGFGGQGPFGGVVGGSSFGGFSGFDDARGSYMSQHQQQQRQNDAMHASRAKEEPGDDMFFQNSLY

na

lP

>OsWRKY25 MPPKRYKHLLLLHRQTQQRRARSRLRLRRPMAVDLMGFSPRGGCRPSVETEQLAFQEAAAAGLRSLELLV SSLSAGGEHHHRRRPQEKQSSPPLGEIADQAVSRFRKVISILDRTGHARFRRGPVVGAAAAEAAAAAASA SPSSSPVSPPLPPVTTQPAAAVKSLTLDFTNPAKVAAASVTSTSFFSSVTAGGDGSVSKGRSLVSSGKPP LAGGVKRKHPHPPCAAAGDGHGHGAGHAHAHGGCHCSKKRKQRVRRTVRVAAASARVADIPADEYSWRKY GQKPIKGSPYPRGYYRCSTVKGCPARKHVERAADDPATLVVTYEGDHRHSPPPPPLV

Jo ur

>OsWRKY26 MAAAAAGASTPFNFCRHGSHAEYDAVFSGSWMARRPSAAPHGGGASGSGSGSGYGAASYVAPTFGAAFRQ QHLDLLDYLSDDQGVPAPPPAAVPSASYVTPAPAMAPAEPVVPDAVAAAGGYPRSVAAAAAAVAGEGRDR TTTDKIAFRTRSDDEILDDGYKWRKYGKKSVKNSPNPRNYYRCSTEGCNVKKRVERDKNDPRYVVTMYEG IHNHVCPGTVYYAAQDAASGRFFVAGISHPDLN >OsWRKY27 MNTFTLFRDEVQKASEKVHRHHANDDEAGLFLSLGLSLGSSPDACQCHASKKDEADAGNGGGDGYLALAL RCAPAAGEPMVHPKRQRATTNSSSSSSICGEYGGGAAAAAVPAGHDDDDRSCMITAASTANRPGRVVLRT RCSAPTVKDGCQWRKYGQKTAKGNPWPRGYYRCTGAPGCPVKKQVQRCNHDTSVLVTTYDGVHNHPITPY AAALPPSSSSSSSAAVAMLASSSSSSTWSELQRAMPAAQSSWSQRNYPIQADVVAKAIWDPKFQATVAAA VASYVRDREQSARVAGGKGAGELFNLAPPC >OsWRKY28 MTRRAGGHATAEFRSLCITSGKKEKKAMGSAWQYIQFMDSWIEQTSLSLDLNVGLPSTARRSSAPAAPIK VLVEENFLSFKKDHEVEALEAELRRASEENKKLTEMLRAVVAKYTELQGQVNDMMSAAAAAAVNAGNHQS STSEGGSVSPSRKRIRSVDSLDDAAHHRKPSPPFVAAAAAAAYASPDQMECTSAAAAAAAKRVVREDCKP KVSKRFVHADPSDLSLVVKDGYQWRKYGQKVTKDNPCPRAYFRCSFAPACPVKKKVQRSADDNTVLVATY EGEHNHAQPPHHDAGSKTAAAAKHSQHQPPPSAAAAVVRQQQEQAAAAGPSTEVAARKNLAEQMAATLTR DPGFKAALVTALSGRILELSPTKN >OsWRKY29

Journal Pre-proof MAGAGDWPFAADEAYADSSAIFAELGWANGLAVVDAVGELLPPLDPPGELATPPPPPLDLPETPAGSSAD GAASSCSTDDADGGKPAAASTEAASKSLTPGKKRARQPRFAFMTKSEIDHLEDGYRWRKYGQKAVKNSPF PRSYYRCTNSKCTVKKRVERSSDDPSVVITTYEGQHSHHTVTFPRAAATAAGFSHIHAMAALAAAPFSAH QQLYSNLQPPPPTMPLAATTPASSSSLLQLPLHCNHELQVVASCGGYPSSSSSPPASVLPVDKGLLDDMV PRAMRHDG >OsWRKY30 MDVYAATSTSTNAIDIGAVASRAIREPRVVVQTTSEVDILDDGYRWRKYGQKVVKGNPNPRSYYKCTHPG CSVRKHVERSSHDLKSVITTYEGKHNHEVPAARNSGHPSSGSAAAPQATNGLLHRRPEPAQGGGGGSLAQ FGYGSAGHRPAEQFGAAAAGFSFGMLPRSIATPAPSPAIAVPAMQGYPGLVLPRGEMKVNLLPQSGNAGA AASQQLMGRLPKQHPQM

ro

of

>OsWRKY31 MDGDAWWYPGGGGGGGSNNWDLGAVVRFGCGGGRVSPAAALLGEAWEYDDDPFSSFLAPPMTAQQAALPA VWEEGDDGDAAWMAPLPGLQTGGGWGDQAPMVVDELCGALVVAPPPPPKQQEVLQVQQQPPPADNTQPTT YQQGSGGDGESTRAGGSRKKQTRKEVVRVAASGPAPDLWAWRKYGQKPIKGSPYPRGYYRCSSNKNCAAR KQVERCRFDPSFLLLTYTGAHSGHDVPLHRNSLAGTTRHKPPPPPPLPSAADKSPATAAEAATASQSPGL SPTTPLRASSMELHGEDDAEAELQVEEDDMAIDDEDDDDVADETISTVPWGTPISDAIIAASYEWR

na

lP

re

-p

>OsWRKY32 MSSKKKRAAIDLSLEAERRRPEERGGGSDREASDGAAAAAEEDGDVKQREGPKEETGGEEEKVVEVVVDQ GEDGSNEEIKYRTQQAEMIEEDKQPAAAANVDDDGGDSDGVGASAEEKHMVTEATGGEGDDGGDSRTPMA QDELSEMQEEMERMKEENRMLRRVVDKTVRDYYELQMKLAAYQQQPAAADEPKETEVFLSLGATAAASAG CGGGFPEAKSKEQAAWRRRSVGSDDSDCGKEDLGLSLSLGASSSYDDDQKAVEARPHDVDGAAAAAMIGG DGSRPAPRGYALLESSKVQGGAAPAAGELAAAGGITSQSVNPANRKTRVSVRVRCQGPTMNDGCQWRKYG QKVAKGNPCPRAYYRCTVAPGCPVRKQVQRCLEDMSILVTTYEGTHNHPLPVGATAMASTTSAAATFMLL SSTTSSSSVSDASAAPSSSYLSPYLLNSASPLLMPGATGGGGGMQHLNLFGNSPSSSSLLAPQAPGSSKY PWSPNHPPLAGAGGNKRPFWSAGGDGDKPAPAALAENVGAVMSDPNKFSAAIAAAINNFMGKDGESSSGK SSTFAWTRSMDCTATPASVPAPARPRREPVHGPGPRRGEQAAKL

Jo ur

>OsWRKY33 MAISLYRPVLRNTSSGCAAAAACADQYSAATPDNSSVTFGDDEADNESHSSEGYEPEAKCWKEDADNEGS SGGMGGGAGGKPVRKPRLVVHTLSDIDVNIDILDAGFRWRKYGQKVVKGNPNPSRGTTPSRCSPIPPAFT AANGAGAAFQRTKDKPRDDLFVESLLC >OsWRKY34 MAAAGEKGKMKIRRKMREPRFCFQTRSEVDVLDDGYKWRKYGQKVVKNSLHPRSYFRCTHSNCRVKKRVE RLSTDCRMVITTYEGRHTHSPCDDNSSGEHTNCFSSF >OsWRKY35 MEEWKDSNHRGADYLMTMPMQNFLADAFPPPELLEGEGGFEKHGLSVAVGSPPPTPPPPEDGCSPLPLTP QFGQKFGSGGGGGGSLADRRARGGFSNVARISVPYNQPAADVSSAGAPSPYVTIPPGLSPTTLLESPVFS NAMGQASPTTGKLHMLGGANDSNPIRFESPRIEEGSGAFSFKPLNLASSHYAAEEKTKSLPNNQHQSLPI SVKTEATSIQTAQDEAAANQLMQPQFNGGKRSRAAPDNGGDGEGQPAEGDAKADSSSGAAAVAVVAAAAA AVAEDGYSWRKYGQKQVKHSEYPRSYYKCTHASCAVKKKVERSHEGHVTEIIYKGTHNHPKPAASRRPPV HPPPPSPATTTTTPLPPGDAQADHAPDGGGGSTPVGAGQAGAEWHNGGVVGGEGLVDATSSPSVPGELCE STASMQVHEGAAAAQLGESPEGVDVTSAVSDEVDRDDKATHVLPLAAAAADGESDELERKRRKLDSCATM DMSTASRAVREPRVVIQTTSEVDILDDGYRWRKYGQKVVKGNPNPSSSSSMDADRSLVVVVVIRSYYKCT HPGCLVRKHVERASHDLKSVITTYEGKHNHEVPAARNSGHPAGSASPGGGAGSSSQPHGVGVGGRRPEVP SVQESLMRLGGGCGAAPFPPHFGLHLPPPPPRDPLAPMSNFPYSLGHAPSPALRGLPPPPPPPPSASALA VAGLGGVVEGLKYPMLAPPSVHSLLRHRQGGGMEAVVVPKAEVKQEAMRPAAAVAGAGRGAAVYQQAMSR VSLGNQL >osWRKY36

Journal Pre-proof MEGSQLETACLPAALYAPLCPYTPPSPPSFLAPLPSLQHKLPQLPQLVHDHAAATGTNHGVMFSSDHGCL YPLLPGIPFCLDSGCGAAACDDDKPAGFAHLGSAEADTSAAAARVDSEIAAAATATTCHGPNSWWKGTEK GKMKVRRKMREPRFCFQTRSDVDVLDDGYKWRKYGQKVVKNSLHPRSYYRCTHNNCRVKKRVERLSEDCR MVITTYEGRHTHTPCSDDATTGAAGDHTASCAFTSF >OsWRKY37 MEGDQAGGDLTDIVRAGGGAMPGSVVVDLPSTAAEWQLPAEPMLFPPPPSLSSTTDGCGAGGAAGADIFG GGGGDLFSGLVDPFSSDYSSGADFLDAMPDAMAKVGFDTAVGGGCGGGGGGGGGSGGHLLDMSRKPLLPR GMPMAAVGGLAAPRVMPSPLSPRAIRPYPPISAGDMMKLGITAGQAAGCAIDAAVAGMQMSSPRSGGIKR RKNQARKVVCIPAPTAAGGRPSGEVVPSDLWAWRKYGQKPIKGSPYPRGYYRCSSSKGCSARKQVERSRT DPNMLVITYTSEHNHPWPTQRNALAGSTRSHHSKNSGGGGGSGSKGSQNDKSQQQPSVKEEQKDQATTAT TTTTSTITTTNSASPVVVKEEEAALAGSSEALELERVMDTTAAGVVDHSELMDHVFSESYKPMIPETGQP DDFFADLAELESDPMSLIFSKEYMEAKPSGGDHAQEKAMAKELDPFDMLDWSTTTNSSAGSSFEQGKRG

ro

of

>OsWRKY38 MGLEDADNEGSSGGMGGGAGGKPVRKPRLVVHTLSDIDINIDILDAGFRWRKYGQKVVKGNPNPRSYYKC TTVGCPVRKHVERALHDTRAVITTYAGAVVQRDPAVGSANGAGAAFQRTKDKPRDDLFVESLLC

lP

re

-p

>OsWRKY39 MEEELCGNNWDLDAVVRLGCCRRRISPAAVAQQVDPFASFLQQGVAMEVAAEKEVGVEAAWSFPELTVRD GGGGGLGRDADELLKAFCAAFPSSSSSKSSPLPTPPPPPPTQPQPEQQKPVTVQENLPAPTTAPARASQP AAARQVPAGGVPRSKRRKNQQKKVVRHVPADGVSADVWAWRKYGQKPIKGSPYPRGYYRCSSSKGCPARK QVERSRSDPNTFILTYTGEHNHSAPTHRNSLAGTTRNKLPSSSSASAASAQPQPPPPSVVVVGAGGGGAE AAGLSPTTPLRTPSMEEDEEEEEEEELLVEDMEMAGEDELLFLNGGDDNAALDGTPMSSLFDIADEPFLP SPWTEPTAAGS

na

>OsWRKY40 MKNSSNKRSLVADQWHPSSVCCDHRAALREIAKGQSLVTQLRAIVLPALHSDERCDLAAQMLEGILDCSR KAVSQLQLLLSSPHDDDDHHHVDDKRRVRKIISSSDDDDHCSSKAAEDHNAKPLRQHKRRRFGDSVSLET PVPHYDGHQWRKYGQKHIKNSKHPRSYYRCTYRQEEKCKATKTVQQREDLHHANSYNGDHPIMYTVVYYG QHTCCKGPAALADDHVVVEASQISTDSHCQSPSSSSDLQAAEVHADGWLDRYYEKIYHCSIA

Jo ur

>OsWRKY41 MFNLPRRLEELLCHHGSMLPKGADEEIPLIKQDLEEIISILHGHILTSTSMPPGLRIDLIFTTIREFSLR IQDALQRHAMYNNLGGVAGTASTTRGDVCSATPWHPTKTQFREHVDNVRSVSIDVDGMEAALNDLNKLKN LLAGIPTASLVQFREHADKVRGIHTDIEAILNKLENIPPGITTTTTTTRGDVSSTSSRQPTLFMESTCHV GIDAAMNKLENLLDLCGEEKLKVVSIVGVGGVGKTTLANKLYRKLRWQFECRAFVRTSQKTDMRRLLINI LLQIRSHQSPDNWKVHSLISSIRTYLQDKRGCVCCLLSLISALATIRPHLAKDIPPIKGFLIVIDDLWAT STWDIIKCALPEGNKSSRILTTTEIEDLALQSCSYDLKFIFKMIPFGEDDSRKLLFSIVFGSHSKCPPEV SETLYDIVRKCGGLPLAIVTVASLLASQLDKLEQWDYINKSLGYSLMANPTLEGMKQLLNLCYNNLPQHL KACMLYLSMYQGDHIIWKDDLVNQWIAEGFICATEEHDKEEISRGYFDELVGRKIIQPVHIDDSGEVLSC VVHHIVLNFVTYKSIEENFIIAIDHSQATIRFADKVRRLSIHFNNVEDAPPPTNMRLFQVRTIAFFGVLK YMPFIMEFRLIKVLFLHFLGDEDSTGIVDLTKISELVRLRYLKVTSNATVKLPTRLQGLPYLETLKIDGK ISEVPTDIYLFASCNSAENLWSLGELSNLRDLQLTYSEIHSDNLKDNMKYLGSILGKLRNLTSITLSPPG SSCPDTLHIDRDTKRRINVDGWSSVSSPPALLQRFELLPCVCVFSNLPNWIGQLGNLCILKIGIREVTSN NIDVLGVLPHKSVDGRETLRRIRVKVSSTVEDGFSWVKYGQKDILGTMYPRSYFRCIHRHTKGCLATKQV QPTDDDHQILDVIYYGEHTCDQSARSDDRQLKSSRPAASSNLQEPQQPGLEQSRPAAKRRRKTVRWKTQV RVSSVQDVGPLDDGYSWRRYGLKDILGAKYPRSYFRCTHRNTQGCVATKQIQRRDGDPLLFDVVYHGDHT CSERASLNEQVTWPRSSASSTEQSSTITYTAAAGSVEDDEEGVTSATNFLSMDDMLDLGGGDVIDMDFPS FDFDAIDAFLLG

>OswRKY42 MADPFPAAARGGEQGGGTAGQLVATPSRLRTAVASMLNRTGHARFRRAAPVVVQEEEDEAAAAARDAVVR CDGLSASASSSFPSSVTGVTGDGSVSNARAVLPAAGAGDKPPPMQSASDYASDGRLKRSSDDDGERCHCS

Journal Pre-proof KKKRKASWRARRRIRVPAISSRNADIPADDYSWRKYGQKPIKGSPYPRGYYKCSTVRGCPARKHVERDPG EPAMLIVTYDGDHRHGEPGHRRPDEAATTTEHRTTDQTTGRLL >OsWRKY43 MSSWAATSAGEMPVAPPSSSSAAVGMVEFPAAAAGLGYAGMTAKEAGGGYQERRVVVGEMDFFKTAEKRG ERKEPPPATATAAASGHAGASPDDLSLNKDDLTINMGLLVGRRRNSGSEESIVDDGGVSSNDEEHREAKA ALAVTKAEIGRLSEENKRLKNMLSNVTTKYNSLQMQFVTLMQQRRSVLAAPIHQQELLDPEKKEQEGSQQ QQQQQLIPRQFISLGSASLQPDVEAPHSVVVVGGGGGDVCAPSSSNPDAAVPAMMPLPHFDHHNHHHPIH GGRERGSSPAEADHHRHRQQEQPPPPPQQQLLPPSWLPADKVPRFLPGKGPEPIPEAATMRKARVSVRAR SDAPMISDGCQWRKYGQKMAKGNPCPRAYYRCTMAAGCPVRKQVQRCAEDRTVLITTYEGNHNHPLPPAA MAMASTTAAAASMLLSGSMPSADGSLMAGSNFLARAVLPCSSTVATISASAPFPTVTLDLTQTAPPPPPA SSTQPQPPRPEPAQLQAALAEAARPVALPQLFGQKLYDQSKLSAVQAVAGTKGSDGGALADTVNAATAAI ASDPNFTAVLAAALTSYIGSSSGSGGGGGGGSSGTVQPLMSGGGDSCSRDDKIGEQNS

-p

ro

of

>OsWRKY44 MAVDLMGFSPRGGCRPSVETEQLAFQEAAAAGLRSLELLVSSLSAGGEQHRRPQQKQSSPPLGEIADQAV SRFRKVISILDRTGHARFRRGPVVGAAAAAASASPSSSPVSPPLPPVTTQPATAVKSLTLDFTNPTKVAA ASVTSTSFFSSVTAGGDGSVSKGRSLVSSGKPPLAGGVKRKHHPNPHPHPPCAAGGDGHGHGAAHAHGGC HCSKKRKQRVRRTVRVAAASARVADIPADEYSWRKYGQKPIKGSPYPRGYYRCSTVKGCPARKHVERAAD DPATLVVTYEGDHRHSPPPPPLPLV

lP

re

>OsWRKY45 MSPAPAPAYAQVMEDMEKGKELAAQLQGLLRDSPEAGRFVDQILHTFSRAMRALDKAAVSAAGGEGSEVQ SEVTCGGGASAGGKRKAPAANRKANCRRRTQQSSGNTVVVKNLDDGQAWRKYGQKEIQNSKHPKAYFRCT HKYDQMCTAQRQVQRCDDDPASYRVTYIGEHTCRDPATAPIIAAHVIHQVAAGDDDDGCGGLHAGSRLIS FVAAPAAPVDAAAAPTTSTITTVTAPGPLLQPLKVEGGIGSSDQEEVLSSLTPGSSAARGGGVAGPFGPD QGDVTSSLHWSYDAVAGMEFFKNDEVVFDLDDIMGLSF

na

>OsWRKY46 RPLGPHKILRFFSETEMALDSVPSYPSDLGSSRARTPQQQRVSPRKEERTWTTDTYAPYDDGHQWRKYGE KKLSNSNFPRCTYKNDMKCPATKQVQQKDTNDPPLFSVTYFNHHTCNSSPKIV

Jo ur

>OsWRKY47 MASPDGGVGDGGAEPHEVMDDLLEMREQAAMLHSMLHGTSPSSCAAAASTRQLNQLIDGVMSRLQSSSLS VMSPGGGGGGRRGSGGRKKKGAKAVAGPHRRSSSGRRRSKSPFVRMVTTKELEDGRQWRKYGQKHIQDSP NNPRSYYRCTHRPDQGCMATKQVQTSESNSSEFVISYYGEHTCRDPSTIPFVVEAEAPAADYANLISFGS SGGASTSRVDPLRQSRHRLMAEAVDPTPSCSFANCHSPVLSSECASEAAALSSSLPLSAVVGSAVTTPST SIVGSAPADYDWPSGLAGGDMAGSFPSSPSSLGFMTGSFGNLPGDDDDMFGFDP >OsWRKY48 MALIATGATATATAAPVASPAASSMASELMAQGRESAAVLEALLHGASLPPAHGGAHALAAEILRCCDRA LAALRAGGDAESSSADTKRKPATAQPSTRRRRRATASGGGAAAAAEPARVEKARTSEDGFLWRKYGQKEI KNSKHPRLYYRCSYKDDHGCTATKQVQQSEEDPSLYVITYFGDHTCSCQTAAAAAMDDDDDDENSQHFVI NFGPATASRSGSPPLLYDDGDDGDVWRETAATPPSSRQSRCSPEGDGEESGVKMSKEEPVDSCPAPSAVS SPADVVSCSSPAMEPDLLGCLNWDDDFGDSSFVDADEFMNFDEIDLFQIYS >OsWRKY49 MSGGGGGGEGFPFHDELASLFAERPPNGAMPGMLQQQQPWSFIDYHHHLMQESAPTTPPLDYEAFAGEFD DDVAPLEEVKRELVVDGVGLFPGGGASAAAAAAAVAGPMTPNSMSVSSTSSEACGVGGGAGGDEESAGKC KKEEEGDGGDDDGKEGSSTTKGDGDGEDKNKKGGKGKGKGEKRPRQPRFAFMTKSEVDHLEDGYRWRKYG QKAVKNSPFPRSYYRCTTQKCPVKKRVERSYQDAAVVITTYEGKHTHPIPATLRGTAHLLGAAAAAHHHG GLQYHHPGHFAAAVGHRLPPQPHDALGGGLLAPPHAQHLHAMQHQMQLAAAAASGGSLHAAAMQQMPQPD HAGLAAIIASTTGASTTPAPPPATGSAAAAATPLRMQHFMAQDYGLLQDMFIPSPFLHNDDANNNNHR

Journal Pre-proof

>OsWRKY50 MEEAYCMMMVGRERELVAELRHLLFPSPSPTPTTPASHSTTALAGDGECCLPPGLTTTTTVSGGGRRRGR KRVNRDNDNVKLLLQADDDQEAVIADHGDANAKPLPNFTKTRRRKQQATTSTMVTTVPDFDGYQWRKYGQ KQIEGAMYPRSYYRCTNSTNQGCLAKKTVQRNGGGGAAGYTVAYISEHTCKSIEPSLPPVILDTTVRTTN NHQQPAAAESPAATSSSSSNMVMTSSETGNWSGQHGAYACRQMIAADEEYCCWDTPATTTTTSGSNG

>OsWRKY51 MITMDLMSGYGRVDEQVAIQEAAAAGLRGMEHLILQLSQTGTSERSPAPAPAQEQQQQQQVDCREITDMT VSKFKKVISMLNRTGHARFRRGPVVAQSSGPAASEPAPVRSSPSAVSRPMTLDFTKAASGYGKDAGFSVS GISAASSSFLSSVTGDGSVSNGRGGGSSSLMLPPPPATSCGKPPLSSAAAAMSAGVGHKRKCHDHAHSEN IAGGKYGSTGGRCHCSKRRKHRVKRTIRVPAISSKVADIPADDFSWRKYGQKPIKGSPFPRGYYKCSTLR GCPARKHVERDPADPSMLIVTYEGEHRHTPSAAGQDHPPAPPPPLALPLA

ro

of

>OsWRKY52 MAVTESACLSYEQEAVAVREVAQVYELIKTQQPLLLVHQQPQQLAHGLLNHALRALNVALSVMNQPHASS SAAAAAVPVMSMIKAEAATPANSSSPAADVAADNHVVGKPRRSSSAAKRRRINGEEYKSSSWSQFTPVPH EDGFQWRKYGEKKIQGTHFTSVNTQEDDGGVAMDS

lP

re

-p

>OsWRKY53 MASSTGGLDHGFTFTPPPFITSFTELLSGGGGDLLGAGGEERSPRGFSRGGARVGGGVPKFKSAQPPSLP LSPPPVSPSSYFAIPPGLSPTELLDSPVLLSSSHILASPTTGAIPAQRYDWKASADLIASQQDDSRGDFS FHTNSDAMAAQPASFPSFKEQEQQVVESSKNGAAAASSNKSGGGGNNKLEDGYNWRKYGQKQVKGSENPR SYYKCTYNGCSMKKKVERSLADGRITQIVYKGAHNHPKPLSTRRNASSCATAAACADDLAAPGAGADQYS AATPENSSVTFGDDEADNASHRSEGDEPEAKRWKEDADNEGSSGGMGGGAGGKPVREPRLVVQTLSDIDI LDDGFRWRKYGQKVVKGNPNPRSYYKCTTVGCPVRKHVERASHDTRAVITTYEGKHNHDVPVGRGGGGGR APAPAPPTSGAIRPSAVAAAQQGPYTLEMLPNPAGLYGGYGAGAGGAAFPRTKDERRDDLFVESLLC

Jo ur

na

>OsWRKY54 MALTMVSLAAIAGKPTAASAQLVAEGRESAARLYALLVGSSALHGPVGLAEQILLCFDRALAKLHGVNLA GAEDDDDAAGNGNGRKRKPGRGRGLTAASAAASSKRMRVSNAGGNGARIERKATMDDKFLWRKYGQKEIK NSKHPRFYYRCSYKDDHGCTATKQVQQSETADDDTASPVYIITYFGEHTCRHGDDAAAMVVDGGEEEDQL SPAQMVISFASSNGGDASVSWPCSGDDAQNNSETSHESSPPEAPAGEEERLRPCTAAGVSDEPIMESTPP APELLADLKPMDGCLLDGESLFGMDELVYFHELSAALGLLDRDWGAPV >OsWRKY55 MRKVDTFAPHNDGHQWRKYGEKKINNCNFPRTNDQCQLTFFKRSTA >OsWRKY56 MSGKGIHILFARSSSHFLQGPYLMGDVLRAQATASAEEVAGGVWPCELDDHLIGELLGDDGLFVPAAEHP TLYYSFGAGSSAAAAAAPCNGGGSADHERRPRPAPAVSRDLCSVYSGPTIRDIEKALSSSASPRPPYPSG RRYSSLYLVEAESKYTSKVRSCGGKMPADGYKWRKYGQKSIKNNPHPRCATRSIIDPI >OsWRKY57 MAHGGGEEEEERVLSHGDVVLLRCDLTILRGPHFLNDRIIAFYLAHLAADHHDDDLLLLPPSVPYLLSNL PDPASVAAVADPLRLASRRLVLLPVNDNPDVSHAEGGSHWTLLVLDNSNAVSGPRFVHHDSLPPTNLPSA RRLAAVLRPLLPASAIPLIEGPTPRQTNGYDCGVFVLAVASAICNWWPTRARHSNSDSDWLEAVKREIKL VHYLVMLDEPSHCPSTQLPSHPSIASTSLPVTAQDFGTWCDMQSHKKVAAVKPVASRPSSRLRSFSMLQE DSTAIDSPRVTSLEEIILRRPKLEDSGTHTAYDQKKADTGKGACWDNLTVSQSVRKPNVSAKNSLSYDGY SWRKYGQKQVKGSEFPRSYYKCTHPTCPVKRKVEMTPDGRIAEIVYNGEHNHPKPHPPRKPTLSTSVETL VATNDAGLENKLEGCDQAIGSDAVVEALRGGCHCLDGFRNGNEISDCKKRPTASLEDTLR

Journal Pre-proof >OsWRKY58 MDGLEAAAGDQQHGRLLIPQLPAAYLASSSMAALSPAGDDWAASLILPDGGSAAAGVGEDDLGGGVMAAA AEESSCGGSSTVTSSGVTEAAPAAATTTRRGRGNGKKAGGGGRTPRFAFHTRSENDILDDGYRWRKYGQK AVKNSDFPSDDELLFSDVDNTQTATENLRFIPLGRVYITG >OsWRKY59 MYMAAAAAGASTPFSFCRHGSHAEYDDAVFSGSWMARRPSAAPHGGGASGSSFGAAFRQQHLDLLDYLSD DQGVPAPPPPAAVPSASYVTPAPAMGPAEPVVPDAVAAAGGGYPRSVAAAAAAVAGEGRDRTMTDKIAFR TRSDDEILDDGYKWRKYGKKSVKNSPNPRNYYRCSTEGCNVKKRVERDKNDPRYVVTTYEGIHNHVCPGT VYYAAQDAASGRFFVAGISHPDLN

of

>OsWRKY60 MNNRGAGGGRGQGDDEEPTPTPPPAPPPETAPSTVGGGGDGVQLVMPEDGYEWKKYGQKFIKNIQKNRSY FRCRDQRCGAKKKVEWHPHDPGLNLRVVYDGAHHHGSPSSAAGEGGTSAAAAANQYDLSTQYFGGAGGPR SQ

re

-p

ro

>OsWRKY61 MERRKVIRMSYTEDDGFSWRKYGQKDVEGAMHPTTQSKSYFRCAHKMTTGCKARKKVQRTDGDPLMVDVV YKGVHSCAGVHSDSQRSSAASSKSNLRPTKSMQVRASSKDVGPPDDGYSWKRYGQKNIFGANYPRCYYRC IHKTTTGCTATKNAQATDGDPLLFDVVYHGEHTCDLQSTHSNDVEPIRPQSGLDDDMCTDDTTTVSTRHD SNTDASSISFQLDWTNCKDESDGPPTTL

na

lP

>OsWRKY62 MEENARLTRALDAILAGHHAHQRALLAPSLSPPPPSATARAPSVSTSCAAREDAAPAVAAAAASTACPSR QQPPTAEPRPKVRTVRVRADAADATDANSMAETVKDGYQWRKYGQKVTRDNPYPRAYFRCAFAPSCPVKK KLQRCAEDRSMLVATYEGEHNHALSTQTTEFVASGCTTSQHAGGSSSSPLPCSISINSSGRTITLDLTNQ AGSGSIASCGVEAAAVSGELVTVLSPELRRHLVEEVVQVLKNDAEFVEAVTNAVAARVVDQIPHIPVHL

Jo ur

>OsWRKY63 MWHVRSNTEDDGLSWSKYEQKEILGAKFPRAYFRCTHWNTKKGCMATKEVQRDDGDPLMFDIVYHGEHTC TQTAESNVDEQIRLTRTRDKKVKRTKRKRQVRVTSVPADDGYSWRKYGQKNVLGFSYLRGYYRCATKGCQ ASKQVQRHDDGLLFDVTYFGEHTCADQPQAAHSSDQSTILERPTFKTRLQGQVDAMEKQHEGFRKMLSSI QEAIGAMSVKQGEVQETMNKMEKSISSWRPQVDAAVQSLQRDMELQRNQVGAVERHQAEADKSSNTSQAL EEREEIARRTSQLPTSPTMSLPVSQALEEREEIARRAPLLPTPPTASMPASTGEIGLDGHRISTQFRGRA SGVVTTLVPPPGKGFVTVTSVKTTDSMDGEDVASRPGPGAGRWHWAPGS >OsWRKY64 MKNSSNKRPLVADQWHPSSVCCDHRAALREIARGQSLVTQLRAIVLPALHSDERGDLAAQMLEGILDCSR KAISELQLQLSSDSPHDDDGHLDDKRRVRKIVSSSSDDDDHSSSKAAEDHNAKPLRQHKRRRFGDSVSLE TPVPHYDGHQWRKYGQKHINNSKHPRSYYRCTYRQEEKCKATKTVQQREDLHHANSYNGDHPVMYTVVYY GQHTCCKGPVASADDHVVVEASQISTDSHCQSPGSSSSELQAAAHAGDSSQCSNISVTCSSSVVVEDCNK LLDMLPAADELTTDVLLFDMTAYAPLDLDINWEMDTNALWA >OsWRKY65 MEEAYCMMMVGRERELVAELRHLLFPSPSPTTPASHSTTALTGDDECLPPGLTTTTTVSGGGGRRRGRKR VRRDNDNLKLLQADDDQEVLAAAVHGDANAKPLPNFTKTSRRKLQTTTSTMVTTVPDFDGYQWRKYGQKQ IEGAMYPRSYYRCTNSTNQGCLAQKTVQRNGGGGAAGYTVAYISEHTCKSIEPSLPPVILDTTVRATNNH HPPAASSSCAAQSPAAAATSSSSDMMMTSTSSTSSETGNWSGQHGAYACRRQMIAADEEYCCWDTPATTT TSGSDGGNSSTCADQVIDLMSGPIRSPLHIAADGNWVDDLLLVGDGLIDISSANITHLFSF >OsWRKY66 MEGEQAAGDLADIVLRAGGAAAAAVAGGGIPSTEWQLPPAEEEEEEPGLFPLPPSSSDGSGMSGADAFGD

Journal Pre-proof PFAGLPDPFGGDYPSSSGGAAAAADFFDAVVAKAGFVDVGVLGGGGGGGCDGGGVDGGGGGSSLLGMSKP ILPRAAMQLPSVSPRAIRPYPVMAGDTVKLGAPMAGGPCAFDGAAAAGLHMSSSPRGAVGGIKRRKNQAR KVVCIPAPAAAGGRTSGEVVPSDLWAWRKYGQKPIKGSPYPRGYYRCSSSKGCSARKQVERSRTDPNMLV ITYTSEHNHPWPTQRNALAGSTRSHHAKNSSSNSSSSGASSASKNNSSHRGYHHHHHHQKPLVKAEPNDQ SAAAAATTTAATVPVKEEAAMVGTSSEALAKTTQKSMDDAAAAASATAAAVEHSDLMQQMFSQSYRPMIP EAAAGGHHDDFFADLAELESDPMSLIFSKEYMATNYKPAGDPAGKEMNAVDKGLDPAYMLDWSSTTVVTR AGGSSFMQGEGGL >OsWRKY67 MAASVGLNPEAFFFSNSYSYSSSPFMASYTPEFSAAAIDANLFSGELDFDCSLPAPAQEYPENENTMMRY ESEEKMRARVNGRIGFRTRSEVEILDDGFKWRKYGKKAVKNSPNPRNYYRCSTEGCNVKKRVERDREDHR YVITTYDGVHNHASPAAAAAALQYAAAAGDYYSPPLSSAGSPPAAYSAGGSLLF

ro

of

>OsWRKY68 MAVDLMGCYAPRRADDQLAIQEAATAGLRSLEMLVSSLSSSSQAAGAHKASPQQQPFGEIADQAVSKFRK VISILDRTGHARFRRGPVESSAPAAPVAAAPPPPPPPPAPVAAALAPTSSQPQTLTLDFTKPNLTMSAAT SVTSTSFFSSVTAGEGSVSKGRSLLSSGKPPLSGHKRKPCAGGHSEATANGGRCHCSKRRKNRVKRTIRV PAISSKIADIPPDEYSWRKYGQKPIKGSPYPRGYYKCSTVRGCPARKHVERATDDPAMLVVTYEGEHRHT PGPLPAPPAAAAVAAMPVSVAVSTGNGHV

lP

re

-p

>OsWRKY69 MEGIVDGSGAQLVVAELVRVQGLLRQLEAHLSAPCSVELCRGLVAQIVALTDRSIGIATRSFSSASGGGA HFADTAPPMPALTSCTPSPLSDGSDHQPFRTTNAKKRKTTARWTSQVRVSAARGAEGPADDGHSWRKYGQ KDILGAKHPRGYYRCTHRNTQGCTATKQVQRTDDDASLFDVVYHGEHTCRPGAASAAAAKRPHAQTLLQS LSASLTVNTDTNTPLTPENRAPAPPLQQQQQQQSVSASPVASDSYGLGGAGYGDWRCCDGDLQEVVSALA TVTSAPDHAAMDAADFMSYCFDFDPAVYGGIVGTPSFFL

Jo ur

na

>OsWRKY70 MTAAPGSLPLVNSRPVSLSLAASRSSFSSLLSGGAGSSLNLMTPPSSLPPSSPSSYFGGVSSSGFLDSPI LLTPSLFPSPTTTGALFSWITTATATAAIAPESQVQGGVKDEQQQYSDFTFLPTASTAPATTMAGATATT SNSFMQDSMLMAPLGGDPYNGEQQQPWSYQEPTMDADTRPAEFTSSAAAGDVAGNGSYSQVAAPAAAGGF RQQSRRSSDDGYNWRKYGQKQMKGSENPRSYYKCTFPGCPTKKKVEQSPDGQVTEIVYKGAHSHPKPPQN GRGRGGSGYALHGGAASDAYSSADALSGTPVATPENSSASFGDDEAVNGVSSSLRVASSVGGGEDLDDDE PDSKRWRRDGGDGEGVSLVAGNRTVREPRVVVQTMSDIDILDDGYRWRKYGQKVVKGNPNPRSYYKCTTA GCPVRKHVERASNDLRAVITTYEGKHNHDVPAARGSAAAALYRATPPPQASNAGMMPTTAQPSSYLQGGG GVLPAGGYGASYGGAPTTTQPANGGGFAALSGRFDDDATGASYSYTSQQQQQPNDAVYYASRAKDEPRDD GIMSFFEQPLLF >OsWRKY71 MDPWISTQPSLSLDLRVGLPATAAVAMVKPKVLVEEDFFHQQPLKKDPEVAALEAELKRMGAENRQLSEM LAAVAAKYEALQSQFSDMVTASANNGGGGGNNQSSTSEGGSVSPSRKRKSESLDDSPPPPPPPHPHAAPH HMHVMPGAAAAGYADQTECTSGEPCKRIREECKPKISKLYVHADPSDLSLVVKDGYQWRKYGQKVTKDNP CPRAYFRCSFAPACPVKKKVQRSAEDNTILVATYEGEHNHGQPPPPLQSAAQNSDGSGKSAGKPPHAPAA APPAPVVPHRQHEPVVVNGEQQAAAASEMIRRNLAEQMAMTLTRDPSFKAALVTALSGRILELSPTKD >OsWRKY72 MENFPILFATQPTSSSTSSSYHFMSSSSGSHDHRHHHGLQAGGNGGGGGGSLSHGLFMGSSSSSIRMEEL SNSQQADDAVVDGGATRSPHGGDGDGAAGDDGGDAQAAAAAGGGGRKKGEKKERRPRFAFQTRSQVDILD DGYRWRKYGQKAVKNNKFPRSYYRCTHQGCNVKKQVQRLSRDETVVVTTYEGTHTHPIEKSNDNFEHILT QMHIYSGLTPSSAAHASSSSPLFPSAAAAASHMFQ >OsWRKY73 MGEVREENERLKTLLSRISHDYRSLQTHFYDVLQQGRAKKLPDSPATDIEEPEFVSLRLGTSTSKCKKED KSTTSSEVKGSTEDFLKIKGGLSLGLSDCRVDANNSEKVQPDVMTLSPEGSFEDARDDTAETTEQWPPSK

Journal Pre-proof MLKNLRSVGAEAEDDIAPQPQVKKARVSVRARCDAPTMNDGCQWRKYGQKIAKGNPCPRAYYRCTVAAGC PVRKQVQRCADDMSILITTYEGTHNHPLSVSATAMASTTSAAASMLISGSSSTSLAAYPAAAASPALAFD ASSKPPLIGGRPFFLPTAAAAAITSTPSYPTITLDLTSPAAAATSSHAAFSLSNRFSHTRYPSTGFTFSG SGPSSAPWPGYLSYGASLSAHPYNAGGGKSSSSFEAALSSINGSRQQGGGGGGGSAPPLYQMQQKAAAAA PPPPSVITDTIAKAITADPSFHTALAAAITSYVGKKGSPPASGGEDSKVGLKWGEHLGLGLTHSSLSTAA AAAASSSSQMFLQPSLGLSGSTTSASTSPVANREQAH >OsWRKy74 MESMEGNGGGRLVVTELSHIKELVRQLEGHLGGSGSPDLCKHLASQIFSVTERSIGMIRSGHFDGHRKRS AAAVAAGDLDSATPSPLSDVSDLPFKATKKRKTSTEKKRHQIRVSSTGGVENPPVDDGHSWRKYGQKEIL GAKHPRGYYRCTHRHSQGCMATKQVQRTDEDAMVFDVIYHGEHTCVHKAVAAGAGKPETETDTNAAAESR LHELSSGLTVKIEGLTAPPQQQQGGGGWNAMPPFCLSSPVSGLAPPDQHNPFSAPSTPENRLAAAASSAA SPATSDSMAAAPFHQAAAAGGDAAWRDAELQEVVSALVAATTATATAQPAPATAMVDADLSALDAFEFDP GFTIDITSFFA

re

-p

ro

of

>OsWRKY75 MHVFAPHQVTPLSVLLAAAAGGGFDRRDRSRVVGGGGGSREGWEMDEVLAQISEGFRLAGELMSELAATQ NDPDYLAARCHAIAGAYAAAARVLQGARHGGGGVGAASTSFGGEGYLHHQHRQQLDLLRMCVGGTAPDEA AAAANPFLGGGGGATTAAFRAPSDAYGGAGAGTSGGPVRRVSSSSRSPPSPVQPRQGGAGSRRRRESGEK VTVMVAAQRTGNTEQPPDDGYTWRKYGQKDILGSRYPRSYYRCTHKNYYGCEAKKKVQRLDDDPFTYEVT YCGNHTCLTSTTPLLTLPAAAATSVTNSPTAAAVLGQDFVMAPAADQQPPQHQQAQPPLSTSIHLGIGWP MTPASLAGAVGEGSTSTTATAPQVTTMGATAAGGGGAAARDADHYPVADLADVMFNSGGSGSSSIMDGIF SSHDRRDN

na

lP

>OsWRKY76 MDAAWRGGVGCSPVCLDLCVGLSPVREPSAARHELLDRPAGCRGGGDSKSMTNDEAKILEAKVTQMSEEN RRLTEVIARLYGGQIARLGLDGSASPPRPVSPLSGKKRSRESMETANSCDANSNRHQGGDADHAESFAAD DGTCRRIKVSRVCRRIDPSDTSLVVKDGYQWRKYGQKVTRDNPSPRAYFRCAFAPSCPVKKKVQRSAEDS SLLVATYEGEHNHPHPSPRAGELPAAVGGAGGSLPCSISINSSGPTITLDLTKNGGAVQVVEAAHPPPPP DLKEVCREVASPEFRTALVEQMASALTSDPKFTGALAAAILQKLPEF

Jo ur

>OsWRKY77 MSSLYPSLLSLSESPAEYRQVGGGRYAGEDVVDDDDDMAAVADAVSSYLSFDMDDVEYYTPEVGFHSKQH NPPPVAAAPLEAGGGREQSRREAAVNLGKMDRGPAPVSGGAATGGVPRSKNGSKIAFKTRSEVDVLDDGY RWRKYGKKMVKNSPNPRNYYRCSSEGCRVKKRVERARDDARFVVTTYDGVHNHPAPLHLRPQLPPPGGYS IAGAPAVVAPHGRLGLEEAEVIALFRGTTATSLLLP

>OsWRKY78 MADSPNPSSGDHPAGVGGSPEKQPPVDRRVAALAAGAAGAGARYKAMSPARLPISREPCLTIPAGFSPSA LLESPVLLTNFKVEPSPTTGTLSMAAIMNKSANPDILPSPRDKTSGSTHEDGGSRDFEFKPHLNSSSQST ASAINDPKKHETSMKNESLNTALSSDDMMIDNIPLCSRESTLAVNISSAPSQLVGMVGLTDSSPAEVGTS ELHQMNSSGNAMQESQPESVAEKSAEDGYNWRKYGQKHVKGSENPRSYYKCTHPNCDVKKLLERSLDGQI TEVVYKGRHNHPKPQPNRRLSAGAVPPIQGEERYDGVATTDDKSSNVLSILGNAVHTAGMIEPVPGSASD DDNDAGGGRPYPGDDAVEDDDLESKRRKMESAAIDAALMGKPNREPRVVVQTVSEVDILDDGYRWRKYGQ KVVKGNPNPRSYYKCTNTGCPVRKHVERASHDPKSVITTYEGKHNHEVPASRNASHEMSTPPMKPVVHPI NSNMQGLGGMMRACEPRTFPNQYSQAAESDTISLDLGVGISPNHSDATNQLQSSVSDQMQYQMQPMGSVY SNMGLPAMAMPTMAGNAASNIYGSREEKPSEGFTFKATPMDHSANLCYSTAGNLVMGP

>OsWRKY79 MALGHHGAARQPPTTMAAAASSSTTSAAAAPATATTTVAFSFQHPTPTPSHHHHHHGVLGYSPLVLDHHH PTTAAASSHAPSPPTLHHHHGGGLHAAAPPPRSSPPHPWSCEEGDHRGRPQMGNKGEAAAAMGAMGINDA GNNTAAAAAAQHHLGVGAVRMKKVGGGGGGGGKARRKVREPRFCFKTMSDVDVLDDGYKWRKYGQKVVKN TQHPRSYYRCTQDNCRVKKRVERLAEDPRMVITTYEGRHVHSPSRDDDDAARASAEMTFIW

Journal Pre-proof >OsWRKY80 MEDWMPPPTPSPRTLMSSFLNEDFVSGSFSNIFSDHESNKPQDQFERNRELVDLSKEVPSQFARPAFQRD ASLDHSLVSPTQRSNSHGGLAERRAARAGFSVPKIDTSRGGSSTVIRSPVAIPPGLSPTTLLESPVFLYN AMAQPSPTTGTLPFLMASNAKSTIPSATKMDEDCTFGNDTFSFQPHVGSRRPNFSAAEKGPNACHQNQSL SNIHQRESSLQSSFTAVKDITDEKNIKTKTSDSMFGDNHSSDEQDDETNQNGENSMPPPNHRSGVPLSHT NDPEVNVLENRGSQTCHNSASLWDNAKNDCLQDVQSEVIETRTAACLPVSTNCDTSIMESQDAVDVSSTL SNEEDDRATHGTASIECNGDGDETDSKRRKLDALTAATAAITTTSNIDMGAAASRGVREPRVVVQTTSEV DILDDGYRWRKYGQKVVKGNPNPRSYYKCTHQGCSVRKHVERASHDLKSVITTYEGKHNHEVPAARNSGH GSSGSGNAPSAPQSNGSQRRQEQGQASFSQFGGAAPFSSFVLPPRNQFGPSASNFPFGMVPPGMAIPMPS LGSLAPAKMAGHPSTMQGYQGLMIPEGEMKTEPMSQLGFPAVNQSSSSFQQMMNRPPSFGPQM

of

>OsWRKY81 MERRKVIRMSYTEDDGFSWRKYGQKDVEGAMHPTTQSNYFRCAHKMTTGCKARKKVQRTDGDPLMVDVVY KGVHSCAGVHSDSQRSSAASSKSNLRPTKSMQVRASSKDVGPPDDGYSWKRYGQKNIFGANYPRCYYRCI HKTTTGCTATKNAQATDGDPLLFDVVYHGEHTCDLQSTHSNDVEPIRPQSGLDDDMYAVVA

lP

re

-p

ro

>OsWRKY82 MERPAPEAAAAAAEPPGAKAEEEEEGPKDVGVKGTGKEKETEKVEAEKEMEGKGKGKESMEEEEGKEKEA KGKESMEVEEGKEKEGKGKAKEKEKETKVKVKEEGGEEKEKGKVEVVEAKRRPAGVGAETPILAVPMVAV PCFLASPAFAGHFSMSHQAALASVTAQAQIQLQSPTTPYSEGLPSPFPITPKAVMPLQRSPSGTEGSVRR SVLEKSASFQSRPHNHVSVNMVGDGFNWRKYGQKQVKSSENSRSYYRCTNSNCLAKKKVEHCPDGRVVEI IYRGTHNHEPPQKTRFVKERVAHITASSGDDETLGLVNNEIIESPSPGCKLEPGAVSEASEQQLFCSSDC EGDAGNKSEDDHPSTEPQPKRSRIIETSTPLTPVLRTVREQKIIVQAGKTSDGYRWRKYGQKIVKGNPNP RSYYRCTHDGCPVRKHVEKAPDDDNNIVVTYEGKHNHDQPFRNNSESKDGPVPMIIPAETTSEQPSTMTS TSEQKQPISLLKDGGDEPMKGKTSEIGGEKAVESAQTLISIKTNPDDMKNTLLKDTSAVVPVQNN

na

>OsWRKY89 MIEHQKALMVELRGMAMPLLPSDNEQAKLALQLLGDILSCSDKAISMLELGGDTKKLTNLVGGKRKGDKH SMDNHNLEEEAKESVSKRRKNAEHTGSTVAQAPHNDGHQWRKYGQKWISRAKHSRSYYRCANSKVQGCPA TKTVQQMDSSGNGTSKLFNVDYYDQHTCRGDGIADPYVVDTAHHSMEPINQNECNSPTLEHEAHEVQDER FENLCMVQNMPEYLIDFELERAFEFIVNSPLGSEHWTFDDSIRCEHSPICIWG

Jo ur

>OsWRKY98 MAAGADERCRALVSGLLSSIDRSISIARSCCTEAAAAGRLTQQAGAAPESPPSADGSAGSDLGADSRCRA NAAGPCKKRKTLPKWSKQVKVRSVQDVGPLDDGFSWRKYGQKDILGAKYPRAYFRCTHRHTQGCHASKQV QRADGDPLLFDVVYHGDHTCAHGVRSAAAAIDGQAAASAEQKHQPTPPQEQNAVSVAFTSMAVVNASTSS PFVSPAMSDCQISYELGGGSMAGVRNVPDVELASKTNSSMGDDMEFMFSLDSDFLDTYKYSSYF >OsWRKY99 MEADDSGGGGGRARRSVEVDFFSDEKKNMKKSRVSGGAAAEADDGKGPAAAGLAIKKEDLTINVSRRRLL RSSIARHLFNSRRSNACARPCSLQLLPAGNNARSDRSMVVDDDAASRPDHEEKSRSSNELAAMQAELGRM NEENQRLRGMLTQVTTSYQALQMHLVALMQQRPQMMQPPTQPEPPPPHQDGKAEGAVVPRQFLDLGPSSG AGGEAAEEPSNSSTEAGSPRRSSSTGNKDQERGDSPDAPSTAAAWLPGRAMAPQMGAAGAAGKSHDQQAQ DANMRKARVSVRARSEAPIIADGCQWRKYGQKMAKGNPCPRAYYRCTMATGCPVRKQVQRCAEDRSILIT TYEGTHNHPLPPAAMAMASTTSAAASMLLSGSMPSADGAAGLMSSNFLARTVLPCSSSMATISASAPFPT VTLDLTHAPPGAPNAVPLNAARPGAPAPQFQVPLPGGGMAPAFAVPPQVLYNQSKFSGLQMSSDSAEAAA AAAQFAQPRPPIGQLPGPLSDTVSAAAAAITADPNFTVALAAAITSIIGGQHAAAAGNSNANNTNTNTTS NTNNTSSNNTTSNNTNSETQ >OsWRKY100 MDGGDIHLLLSILADGEEQARQLGEPAAAADDEYHGGGRGEEYYRGVARQLQGTLARAMGIARAIEAAAF AGGGGGGGASGSRGTTGDRSDSPRSADESSGRTARDAAVAQQERHHDTIKRRKGLPRWTEKFRVPDASLE ATPDDGFSWRKYGQKDILGAKFPRGYYRCTYRNAQGCPATKQVQRSDADLAVFDVTYQGAHTCHQKQRRA AAAGDQPPPPPPQADPSVELLVNFRHGLKVETNGLAPPPPPPPTTTTNFHDDQHFCFPSMPPFHAGVGPP

Journal Pre-proof PPPDDALGGGCNNFSSPPFVSPAGSAAGESYFSMEHSYEPRGGGGHFVMSRGDSSELHEVVSAAASSSAV VDPAAAAGGFDYPLYHGEVDPHLPFPPLFGHASMYGQYRDA >ChrWRKY1 MDTSIPFPRPINARGPAPGQTPSQLSSLPPSLQARLGLGATHDSPVLLPLLQQVEASPTTGIHQLCPPLF QPAQPARVPLPIPARTEAASAAPEPTRAIKREYEPRAGNGKQSVANSDGWQWRKYGEKLVKGSPNPRSYY KCSHPGCLAKKIVERSDSDGTVLSTEYKGDHCHPAPSAMLPIPEALKSDFPVPHAAGAAAAHEDDTDTSE PEPAAALKAAPQDTRAAQAAATAIRKVRDSAESPSKRLDMLAAYAEEAERQLKSSSNSPEQGPSAKRQRT EAGAMRTRANPDDDDDGSGAPSTSGMQRVVDITNMDDGYRWRKYGQKQVKGSPFPRAYYKCTHMGCSVRK HVERSAEDETRFVVTYEGTHSHRLPTGSRRRSARDMAEDDEDYEGEDAEEDSSQPTSPQYGNVNGSGGPG QHAASKAAAQGAQLVHPSGAQPASADFGQQLQQLSTSLLASTVLQQAALSGVLPLLQYNSLSSEALASLG VNSEALQGVEQLNLASVGNLADLTNLLRQHAQMDLALAAQAQAIDAANANWDPLACLITPRPNVSPAGQG HAMGQAPSAGTGRQTKAAVFQKQVATTEA

-p

ro

of

>GhWRKY1 MNSSSSSWVDSLDLNASSTKSLQLETPVSHPNTSNSLIVFGRKLSVKEESGALMEELNRVNAENKKLTEM LKAMCESYNALQSQLVDLMNKNTEKELSPTKKRKSETSNNNNGNIIGNSESSSTDEEESCKKPREEIIKA KISRAYVRTELSDTSLVVKDGYQWRKYGQKVTRDNPCPRAYFKCSFAPSCPVKKKVQRSVDDQSVLVATY EGEHNHLPPSQIEATSGSSRLGSVPVGSTPVKSSGPTITLDLTNSIKSSDEARNSKPKLDSPEATQYLVE HMASSLTKDPNFTAALAAAISGRMFSTNSN

lP

re

>GhWRKY2 MENYQMFFPISAPSTAAQSLPLNMAPNSQAFNSFHGNSVDGFLGLKSTEDLIQKPEAKDFMKSSQKMEKK IRKPRYAFQTRSQVDILDDGYRWRKYGQKAVKNNKFPRSYYRCTHEGCKVKKQVQRLTKDESVVVTTYEG MHTHPIQKPTDNFEHILSQMQIYTPF

Jo ur

na

>GhWRKY3 MAEKRTRSKLSAPTRPSITLPPKPAMDGLFSSGLGLSPGPMTLVSSFFSDLDSTENRSFSQLLASAMTSP GARLPQASMDGSLMEVGFKDGGEKSPGFKHNRPLNFDSPLFTAPTGLTPSGLLNSPSFFCLSPQSPFGIS HRQALAQVTAQAALAQSQVHPQTEYQALTVSAPSESLIRHPSFTPEETSQLMPPLASEPQSSAIEYSEAS QSDRKNQLPIAVDKPAEDGYNWRKYGQKPIKGCEYPRSYYKCTYLNCPVKKKVERSAYGQITEIIYKGAH NHEKPQQNKQAKGGSDGNANSQANPEFGSQVAALNSNKLSETMSSHSILGKNSESTQAAELPGSSDSEEG CDEESREERDDDEPNLKRKNTAPEAAVVLSHKTVTEPKIIVQTRSEVDLLDDGYRWRKYGQKVVKGNPHP RSYYKCTSAGCNVRKHVERASTDPKAVITTYEGKHNHDVPAARNSSHNMVNNSLSQTKPHKVVAENQNHS LLKAMDFGNNGQGPAVLRLKEEQIRV >GhWRKY4 MDDASTPNRSPLPPTIQFPVNPNCTNDEIPRRVIGEMDFFAEKNHKLVDDHVDDSDHANATDINGSIDRT ALELNVNTGLNLLTTNTSSDQSSVDDGISSNMEDKRAKNELAVLQVELERMNTENQQLREMLSQVTSNYN TVQKHLVTLMQQQQDGKPEKNDEQKKSNGGLIVPRQFMDLGIAAAAAAEVDTDEPSLSSTEGRSHDLSGS PNVEVASKEFRSRKRGSSEEGRGIGREDSPDGNKVPRFNSSKTVDQTEATMRKARVSVRARSETPMITDG CQWRKYGQKMAKGNPCPRAYYRCTMAAGCPVRKQVQRCAEDRTILITTYEGNHNHPLPPAAMAMASTTSS AARMLLSGSMSSADGLMNSNFLTRTLLPCSSSMATISASAPFPTVTLDLTQIPNPLQIPRTEGQFQVPFP NPPPHNLPNSQLSPLPLLPQIFGQALYNQSKFSGLQMSHDIEQPQMSHQHQQGQQKSLADTVNAATAAIA ADPNFTAALAAAITSIIGSSHNNNVANNNGANLTSATNSNGNVTTTTTTTTSNSNSNGNNKISNSSNFAA N >GhWRKY5 MEKTMGWEQNTLLNELAQGRDFTNMLRKHLHPSSSPETRQVLLDKILCSYDKALSLLNCSRFMVETKPRV RTLGSPENDASDNKDMFKKRKTSSGWSEQIRVCSAMSLEGPLDDGYCWRKYGQKDILGSNFPRAYYRCTH RYSQGCLAGKQVQRSDEDPTIFEVKYRGRHACNQVPHLVATPKEKGNHYREKQQVEEKQKQSKEMLLSFE TGLKVKTEDLDNREDIFPSFSFPIESEEVQNGLLLNSLMKNMSPAFVSPATSESNYFSVSAFHMGNFDFG QNVQTSESELTEIISAPASVTNSPIVDLDISSLEKLELDQSFPYDNPEFFTNFLQ

Journal Pre-proof >GhWRKY6 MEKQHSRELTFLHSGDSLRQNSGVADRTLDDSRDHVEPHIKEMDFFSSQYQSHDHHHHHQEGNTNNNGSS SFIDSSVNIGLNLLSSSPGVSRATNEEKPKAQESALKMELEKLNEENRRLRSMLDQITKNYNELQGQLLM ALQKQAHRNQQDQKDAVNGMASTIMSVQQFMDPRPSAALDVNEPSGSDDKTQELSASPENTIEIVSKEYD HRMVQNMPGKQVSIEDGTDQTSQSWGSLKSPKIDQSKNEDQVSEGPFRKARVSVRARSEAPLISDGCQWR KYGQKMAKGNPCPRAYYRCTMAVGCPVRKQVQRCADDKSILITTYEGNHNHPLPPAATAMANTTSAAAAM LLSGSTKSKDGLPTSAYYPSLPYASSMATLSASAPFPTITLDLTQGPNAVQLLRPPPSGASFPLPLHGYP QLLGNPMFGPPKLSTASPAMQLGQRPASMVETVTAAIASDPNFTAALAAAISTIMGPPRNQGNSSNNNDE NNNSSNGVGAPALPGSPQIQQSCTTFSSKACTNKPG

of

>GhWRKY7 MGSVSSWPEGVPTSNKERVIKELVNGQECAKQLQILLHNWCEKNGRLSAEELLHKIFASFDHALSLLTSV ESAEDSQNQATSYDDSPCCNGRSEDSTNSRKKLPSKEKRGCYKRRRDEHARTVVSSTVEDGHAWRKYGQK QILNSKHPRSYFRCTRKYDQGCKATKQVQRLEDAGSQMFQTTYIGTHTCRPDSFKAPQIIIDSEPRESYN KFSYGSSNPKIPTKKLHEITPPVKQETKEETPTTSGLTDMDSVVMWKDIIGAEYGDVVSNVYTSTEITSQ NLELDLVMKPVEFEDDFQFDESELV

re

-p

ro

>GhWRKY8 MAKTSGLSIDFDPNYFLIHNPPVLNSFPEDTPEDTNKNNNNNNTMFSGTTTTPENMSLRSTVDRFPVKFD CFYDQTRPSFLPSDDDKRRVVGELDFFAEKNDKLVDDADFMHTSDAGLELNVNIGLNLLTTNTSADQSTM EDKKANNEVAVLRAKLVQMNAENERLKEMLIQATENYNTMKIHLCSLMMKNGGKAKEEDEENKKRNGGII VPRQFIDLGLAVAADADESSSSSPEGKRQREDSSGGDNKVPRLDVDQSEAAMRKARVSVRARSEAPMITD GCQWRKYGQKMAKGNPCPRAYYRCTMATGCPVRKQVQRCAEDRTILITTYEGNHNPPLPPSAMSMASTTS SAGRMLVSGSTSSPDGLMNSNLLTTTLLPCSSTMATVSASTPFPSVTLDLTQSSFSTPPPPYNVANSVAV TAPLVPQVSGQALDNLSKFAGIQMSKDLDRLQQGEQNSIGDTVNAATAAIATDPSFAAALAAAITSIIES SQSHPNNGHNSNAM

na

lP

>GhWRKY9 MENYSIVFSDGSSSSPSSLTMANHHHQQRSYLQSNNGVNTCKNMGASIDKNEGEVAGKKGDKKEVMKKHK YAFQTRSHVDILDDGYRWRKYGQKTVKNSKFPRSYYRCTHKDCNVKKQVQRSSKDDEIVVTTYEGIHTHP VEKLFENFEHILKQIQTYNPL

Jo ur

>GhWRKY10 MAETESQAKRHEGKQYDNSEGEEEEEEEEEEEEEVDELEDEEEEEEEGVSRVNESHGAGTAPSSDAEVRD SQLETLTNDDRCERLQENPTSELKEGAESKEQVKIAHQKALPSLAVQRAEAPKQHLLQSSVSPTSSSQPS PTSITQDISSPPTPTLPAQSHSDQKVNGASPPVANQQNPSNLKMLSFVPVVKTPVSDGYNWRKYGQKQVK SPKGSRSYYKCTFSNCQVKKIECSDQTGHVIEIVNKGIHSHEPPRKMNFTRENKILSSAVPLSIVREQPS RIPNDSDPSTSSKESLPEPTVNTERKRQCSSGSDGHGDVQMKKEFLSEAEPNKRRKKGDAVCPGSVVKTG KKPKFVVHAAGDVGISGDGYRWRKYGQKMVKGNSNPRNYYRCTSAGCPVRKHIETAVDNTNAVVITYKGV HDHDMPVPKKRHGPSSAPLVAAAAPASMNNLQLAIADGEQKQVTSTKWSVDTEGELTGEAVDLGGEKAME SARTLLSIGFEIKPC >GhWRKY11 MGSSAWVDTTLDLNINPFHHRNKALRTKSEGEMADSDVKLPFKQEQNADLVEELNRIKAENKKLTEMLTA LGHCYNDLQNKFTELVTINSENEVSTSKKRKAECEDYSTNMIGFSNGNKETSCSDEDSCKVPRECIKTTI SRIYVRTNPSDNSLIVRDGYQWRKYGQKVTKDNPSPRAYFKCSFAPSCPVKKKVQRSAEDPSVLVATYEG EHNHARPSPSEVAILSPKCSAKPVSCASTRPSAPTATLELMQPEGYGDGAKKKIQEVDDAPAAIQQVLVQ HMAASLTRNPNFTAALAAAISGRVI >GhWRKY12 MAVDLMSFPKMDDQMAIQEAATQGMKSMEHLISLLSHQSNRLDCAEVADLTVSKFKKVISLINRTGHARF RRAPIQSSSSFSSPSSSAASLTVTDSHKALTRTPAPVEAPPLTVNPAPVEAPSTAPVSFVPSQPQSLTLD FTKPCLFSSNTKSTELEFTKESFSVSSNSSFMSSAITGDGSVSNGKQGSSLFLTPAAAVSAGKPPLSSAP FKKRCREHEHSENVSGNGKCHCSKRRKNRVKKVIRVPAISSKIADIPADEYSWRKYGQKPIKGSPYPRGY YKCSTVRGCPARKHVERATDDPSMLIVTYEGEHRHGEGAAQENMAPGVGLVFKST

Journal Pre-proof >GhWRKY13 MEHRGNREQGTLISELIQGMELAKQLRVHLDTASSAESMDLLVQKILSSYEKALLILQFSRPMSMGQPQQ NVGATSGVPESPLSINGSPRSDDLDKDNQDTRDVSKKRKMLPRWTDHVRVTSENLLEGTHDDGYSWRKYG QKDILGAKYPRSYYRCTYRHTQDCWATKQVQKSDEDPTIFEITYRGTHTCVHGNQPVPPPASPEKQEYKL TNLNTNNNQQLQSQAGTLGGRREASQR >GhWRKY14 MAGSSNELSLSVPFHTATSVASAPVEVDVDELNQSGNSNSGMQSGQTDHRVSGSSTISDDGYNWRKYGQK HVKGSEFPRSYYKCTHPNCEVKKLFERSHDGQITEIIYKGTHDHPKPQPSRRYSSGNTMSVQEERFDKVP PLTGRDDKSSNMYGQMAHSIEPNNTADSSPVTANDDNVDDVDDDDPFSKRRKMDGGIDIIPVVKPIREPR VVVQTLSEVDILDDGYRWRKYGQKVVRGNPNPRSYYKCTSAGCPVRKHIERASHDPKAVITTYEGKHNHD VPTAKTSSHDIAGPVTVNALSRIRSEENDAISLDLGVGISPAPENSSNEHQQLHSELLQSHHQTGGSSFK LHQVTPMTAYYGVPNGGMNQYGSREAPSEGRTVDITPLNHSYAYSQTMGRILTGP

ro

of

>GhWRKY15 MENFQGFLTTSSLLGYGEFQSGKGLMEEMEGGAENKTFAGLLGTQKKKGDRKKIRKPRYAFQTRSKVDIL DDGYRWRKYGQKAVKNNKFPRSYYRCTHQGCNVKKQIQRLTKDEGMVVTTYEGMHSHPIQNSNDNFDHIL SQMQIYTTSF

lP

re

-p

>GhWRKY16 MEPVDHHIVSLVQSSDGIKRVVNEMDFFADDRSSKQHDKADDDVHHHHHVKTESEHHGADQDDDVNTGLN LLTTNTASEKSSVAGDGITSSQNLKEKQRVNQLADVRAELDRINAENQRLKLTLNQVNSNYYALQMHLVS LTQRHRNRIAESSEANTQMNRPIEEKKHGEGMVARQFMDLVQTETKAEKDELSESSSEDRCHELSGSPGN TIVESMERRQKNNKSRNQREDTPEGLPHPAGWLPNKGPKFNTSKDNEQAQQTMAMIRKARVSVRARSEAS MISDGCQWRKYGQKMAKGNPCPRAYYRCTMATGCPVRKQVQRCAEDRTILVTTYEGNHNHPLPPAAMAMA STTSAAASTLLSGSMPSADHGIMNSNIVPKGMLPGSPNLITLSASAPFPTVTLDLTHPPNQRPLHQLHPL SPNWPHNISRLPAHLLGHPIYNQSKLLGTYLSSQGNNELPVHLSQNQMQGHPMVDTAATAAAITADPNFT AALVAAITSIIGNSHRDNSGNNNSTTSRNTGDNNT

Jo ur

na

>GhWRKY17 MEEQILASSSSSSSSSSAVAAATMQHDLSGSAGSGGARYKLVAPAKLPISRSACITIPPGLSPSSFLESP VLLSDVKAEPSPTTGSLIKPQAVHASVACSTYSATTACSNAFDERNPSTFEFRPHPRSNMAPADLDPQRS EQSRQIQIQYQTISFNSSASVKSEMAGSSNVLSLSVPVHTATSMASAHAEVDAEELNQIGNPNGGMPSGQ SDHRGVGPVSSEDGYNWRKYGQKHVKGCEFPRSYYKCTHPNCEVKKLFERSHDGQITEIIYKGTHDHPKP QPSRRYSSGNIMSGQEERSDKVSSLNGRDDKSYCGYGQMAHSIEPNSTADISPVTANDDNLDEVEDDDPF SKRRKMDGGIDITPVVKPIREPRVVVQTVSEVDIMDDGYRWRKYGQKVVRGNPNPRSYYKCTNAGCPVRK HVERASHDPKAVITTYEGKHNHDVPTAKTGSHDTAGPVAMCGPSRVRSEENDAISLDLGVGISSVPEHAS NEQQQLDSELVQSRPQAGGSSFKFVQPTMMAYYGALNGSMNQYVSRENPSEGRGVEITPLNHSYAFPQSM GRILTGP >GhWRKY18 MASSYDTSANSLPQNAFGFSTHPFMTTSFSDLLASGTTDDDNSSSAVNHGKRGGGLSLSDRIAERTGSGV PKFKSLPPPSLPISPPPVSPSSYFAIPAGLSPAELLDSPVLLNSSNILPSPTTGTFPAQAFNWKPSSGNN QQNVKRVDKSYTDFSFQTFQSSTNTNQTAQQEAWSFKESVKEADFSSGKNMVKSEYNQMQSFSPEIANIQ SCQSDYTNYQHQQSQSLRGNRKSDDGYNWRKYGQKQVKGSENPRSYYKCTYPNCPTKKKVERSLDGQITE IVYKGSHNHPKPQSTRRSSSSSSSSIAIQASSAATSEIPDHSFISHGSGHMDSAATPENSSISVGDDDFD RGSQKSKSGGDEFDEDEPDAKRWRGEGETEGISAPGSRTVREPRVVVQTTSDIDILDDGYRWRKYGQKVV KGNPNPRSYYKCTHPGCPVRKHVERASHDLKAVITTYEGKHNHDVPAARGSGGHSVNRPLPDNNNNNGAM AIRPSPLNNPMSNNQALRGLRQSTTEGESPFMLEMFQGNGNLGFSGFGNSGGSIMNQQHDVHNMFSRTKE EPRDEAFIESLLY >GhWRKY19 MENWDLQAVVGGNSSNDHPTENPEFSFGPWSFQQDEDFVRFPGIFETNPKVLDELEQLYKPFYPDLNPFS TQTIITSSIPVPLHVDEPAEKRKKKQSFTVSQSDISASPKPRRFRKNQQNRVVEHVTADDLPSDVWAWRK YGQKPIKGSPFPRSYYRCSSSKGCLARKQVERSSSDPRVFIISYTAEHCHGHPTRRSSLAGSTRSKPLTA

Journal Pre-proof AKSIEAHAEGETVKQERMKMEVQLHGEEEGGKILSPDLLLSDDELIRRLENFDEGFFVDQFPHFSHEM >GhWRKY20 MHNKLPSLQNSIKDQKSRFSLKMDFSHQNPNSNPNYTFFPENVNPMPDFEVSDYLDLDGSIFDDDTSSEK GMAAATELSGATSKNSNRKCKNGVGKNKSELGVRVAFRMKSEMEVTDDGYKWRKYGKKSIKNSPNPRNYY KCSSVGCNVKKRIERDVKDQSYVVTTYEGVHNHETPSMVYYNQMSIMAPNAWTFQASPPSSSST

of

>GhWRKY21 MSQQAMLNQELFEEKEMSQMGFFTFPQTLTSYPPPFGCLKAFNIAPPPANLTETLFSHPKHREEDSASDF AGPQLLSLHISTPNLWAWGEVNECLGTKKSSGDDHLGVSAMKMKRIKGRRKVREPRFCFKTMSDVDVLDD GYKWRKYGQKVVKNTQHPRSYYRCTQDNCRVKKRVERQAEDPRMVITTYEGRHAHSPAQDSEDSHHPYSH LNNFF

re

-p

ro

>GhWRKY22 MAGLSNELSLSVPVHTATTMVSVPPEVDVEELSQMGNPNIGIQSGQSDHRVGGQSVPFDDGYNWRKYGQK HVKGSEFPRSYYKCTHPNCDVKKLFERSHDGQIMEIIYKGTHDHPKPQPSCRYSSSNIVSGQEERSDKLS SLNGRDSIYGQAVHSVEPNSTADLLPVTANDDNVDDVDDDDPLSKRRKMDGAIDIIPVVRPIREPRVVVQ TLSEVDILDDGYRWRKYGPKVVRGNPNPRSYYKCTNAGCPVRKHVERASHDPKAVITTYEGKHNHNVPTA RTNSHDTTGPIPVNGPSRIRSEDNGAISLDLGVGISSISENSSNEHQQLHSELVQCHPQTGGSSFKFVQP HPMAAYYSVLNSSDMNQYGSRENPNQGPGVEITLLNHSYAFPQSIGRILMGP

na

lP

>GhWRKY23 MDMEGVEEANKAAIESCNKVLSHLSLSKDQIQFSNLMIETDEAVFKFKKVVSLLSSNGRKRRKFKKITSN LPENIFLETPNCTTTLSPKPLQVYPHPNLLDKRTSIEVDFRAQHQLQLTQNLFDPNNQNPSFQMMNSSLV HHHHHHHQQRVQFQQMKNQAADMMFSKSNSGMNLKFDRSLSPTMSFLSSLSMDHGSVNSLDGNSFHLIDI PRNFGQLSHQSGRCSFRGEDGNFKCGTSSTKCHCSKRRKLRINRTIKVPAISNKVADIPPYQYSWRKYGQ KPIKGSPHPRGYYKCSSVRGCPARKHVERCLEDPSMLIVTYEGDHNHSGFLSTQSAHT

Jo ur

>GhWRKY24 MAGIDDNISIIGDWIPPTQSPRTFFTVMLGDDSGSKQVSKAPLENKSEGLFLGENSDKMDVNQGCEPSNE STFEQKSSTRAGLLERMAARAGFNAPRLNTENIRSTDTSLNPEIRSPYLTIPPGLSPTTLLDSPVFVSNS LAQPSPTTGKFSFIPNVNDRSCIPESRDKSEDNYFEDINASSFAFKPIAESNSSFLLGATSKVTPASLPQ QSFSNVEVSVQSENSQPSRSVDPRKLQQQNSNMLSLQADFLRSSTEKDTGSNNSADQRVFDPVGGSVEHS LPLDEPQDEEGDQRFSGDCMAGGAPSEDGYNWRKYGQKQVKGSEYPRSYYKCTHPTCQVKKKVERSHEGH ITEIIYKGAHNHPKPPPNRRSSIGSSNPLIDVQLELPEQTGLQNGTDGEPVWATAQKPMVAATHDWTHEN IEMTSSATICPENGTAPAPNGTHIESGDAVDASSTFSNDEDEDDRGTHGSVSLGYDGEGDESESKRRKIE AYTTEMSGATRAIREPRVVVQTTSEVDILDDGYRWRKYGQKVVKGNPNPRSYYKCTSAGCTVRKHVERAS HDLKSVITTYEGKHNHDVPAARNSSHVNSATSNTATALASSSVQTHAHRSEPSQLHNSMVRPTSFGPFTL PGRQQLGPSPGFSFGMNQPGLANLAMAGLGPGQPKMPVLPLHPYMPHQRQGNEMGFRLPKGEAKMEPMSD PGLDLSNNTSVYQQLMSRLPLGPQM >GhWRKY25 MEGERGVPNYMLQVSFSSSTPPPQAIHETGFLQFEENQVMSFLSPSQPLTTSPVSNTTNSTTALRFITHN DQVGTLDPIPKAVHDENCGSNVKDGNNNSWWRSSALEKNKVKVRRKLREPRFCFQTRSDVDVLDDGYKWR KYGQKVVKNSLHPRSYYRCTHNNCRVKKRVERLSEDCRMVITTYEGYDLLLVSIAAFYVLMVPYTKVEES FNIQAMHDILYHRHHLDNYDHLEFPGVVPRTFIGAFLVSIFSSPIVLAVQLLHLPKIYSLIIVRLALGCI ILSTLRFLRIQVRNKFGHQVEAFFVIFTAIQFHLLFYCSRALPNILAMGVVNLAYGHWLKGNFYTALNYL VFATTIFRCDVVLLLCPLGLELLLTKSISFWRAFKCCTVTTLLCIGLTVLVDSIMWKRFLWPEFEVFWFN SVLNRSSEWGTHSIHWYFTSALPRSLLTAYPLFMLGVLLDRRLLPLVLPALSFVVLYSKLPHKELRFIIS SVPIFNLSAAVAASRIYNNRKKSFWKFVNLVMLGLLLISLGCTIITFMASYENYPSGRALKELHQIGHLA NKSNELWVHIDPFSAMNGISLFCENEWPWRYSKEEEITLEEFGQRNFTYLINEHRTIDGYKCLFYVNGFS RLRRQSGFPPIILDKEPKVYIHGNIKNEELMLKPWPGCS

Journal Pre-proof >GhWRKY26 MDVDWDLHAVVRGCATVTTTTSSSGGGCGGVAANSSLMADLYPQSRFSSFGSQEEPFQGQFSCFPNQFEA KNAMEELHKLYKPFFPKSQPQTKQQQEHQQVKPKLSQVGSVTTTSSSTSTNNSRSKRRKNQLKRVCHVPP EGLSSDVWAWRKYGQKPIKGSPYPRGYYRCSSSKGCLARKQVERHRSDPNMFIVTYTAEHNHPAPTHRNS LAGSTRQKPFTPQTVTAGDSVKPSSSKPADSLSPTASVDEGVVVQSTKVDSREDLVEDEGEDEFGMSDTA VSDDFFEGLEGLAEIVTRNCFPDNFPASFDFPWTANNAATAAGGI >GhWRKY27 MDMEGVEEANKAAIESCNKVLSHLSLSKDQIQFSNLMIETDEAVFKFKKVVSLLSSNGRKRRKFKKITSN LPENIFLETPNCTTTLSPKPLQVYPHPNLLDKRTSIEVDFRAQHQLQLTQNLFDPNNQNPSFQMMNSSLV HHHHHHHQQRVQFQQMKNQAADMMFSKSNSGMNLKFDRSLSPTMSFLSSLSMDHGSVNSLDGNSFHLIDI PRNFGQLSHQSGRCSFRGEDGNFKCGTSSTKCHCSKRRKLRINRTIKVPAISNKVADIPPYQYSWRKYGQ KPIKGSPHPRGYYKCSSVRGCPARKHVERCLEDPSMLIVTYEGDHNHSGFLSTQSAHT

-p

ro

of

>GhWRKY28 MSPRWDLKVHEVAESSFRRANHLFSCICDGSQKRSNQEISLMAQDTVNGFRKLLSLLDGSMQSECKRIRK GPLPKSHNVNPAELMDSPSFASQSSICKPSQPHILGQFVSPQFNQSNTGLIPRNVVVLPNLIMGLQHSST LPTMMPMFDKKLIPWSSSEIVVPQDESCMLLSKRKIGVGSEEASTKCAASTGGCHCSKRRKLRIKKTIRV PAISNKPSDIPPDDHSWRKYGQKPIKGSPYPRSYYKCSSTRGCPARKHVERCLEDPSMLVVTYEGDHKHS RITCQFPSNVIV

re

>GhWRKY29 MVVGVPSISLEGLVLWIKVPDTRRSALVGVKMGVSNVEVVVNAIAPKRGENQYRIMLLILVLVIAFLKWL LFVGRKHRVKRSIKVPAISNKLADIPPDDYSWRKYGQKPIKGSPHPR

na

lP

>GhWRKY30 MEEVEEANKAAVESCHRVLSSLSQPKDQIQYRNLMAETGEAVFRFKRVVSLLNTGFGHARVRKLRRLDTP FPQSILMDNPHHKVIHESSVKELSSHAKNCLYLGNQHQTAQPASSGHYRILQQHQQQQQLRQRLQLQQQQ MKHQAEMMFRKSNSGISLNFDNSSCTPTMSSTRSFISSLSMDGSIANGGGSFHFNGGSRSSDQGSQHKKK CSGRGDDGSVKCGSNGKCHCSKKRKHRVKRSIKVPAISNKLADIPPDDYSWRKYGQKPIKGSPHPRGYYK CSSMRGCPARKHVERCLEEPSMLIVTYEGEHNHPRLPSQSTTT

Jo ur

>GhWRKY31 MSWNTKKAIEELVRGKELTKQLRDLLTKSLTDDGFVNSEDLVLKICDTFANTLSLLTNSNTTTTTAVADY YYNDDFNDVSQKPTINITSDGRKSEDSGDSIKSPPSTHRDRRGSYKRRKCVDSRKKITSALVDDGYAWKK YGQKQILNANHPRSYFRCTHKTDQGCQATKQVQKLEDDPPKYETIYYGHHTCKNMLKSCHFMVEDSTCNN DSSILLSFANNSTLTNKQGNDNPFFSSVKQESKEDYKPISEITYNGSSSSEYLLSPHHHQLSTFDLSSAD MTVLSPDHDVLSDVVDSVDLDDLLQF >GhWRKY32 MANMRFSDEIPSAFGGGGGGSIFDMAGTSCEGSGGDKWGSLGFMDLLGIHQDFVAPSLFDSFQPSILPPS SPPLPVFHHHDDDILHEPHTDSKQQHLNHQAGLLSPASTVPESSEVLNNPATPNSSSISSSSNELAANDE QTKAGNDEDKTKKPLKPQKKNPKRQREPRFAFMTKSDIDHLDDGYRWRKYGQKAVKNSPYPRSYYRCTTA GCGVKKRVERSSDDPTIVVTTYEGQHTHPCPIMPRGAIGIAPDSSAFGAPSFIIPQPMYINHQQQQQQQL QPYIYNSTPSLNITTASSNIIHPSFPGFLQEKPRFSNPNPSALSHRDHGLLQDIVPSPMRTEAKEEDQ >GhWRKY33 MEDKEGVDPASEFTSNSSWALAGGPDSVSDSVNYFFDKESNILSEFDWNLQPDHAEQIDRFDELDRADTR PYLAGSFSGSHSCVAAAACGSVSGSGTASSPAGGAADVSTLIPSVSSSSGEDPSEKYTGSGGKPPEIPSK ARKKGQKRIRQPRFAFMTKSEVEHLEDGYRWRKYGQKAVKNSPFPRSYYRCTNSKCTVKKRVERSSEDPT IVITTYEGQHCHHSVGFPRGGLISREAAFAGRFTPSVSQFYYPHGEIPASIIRSQQVPIEIVESRPLPQT TNEGLLGDIVPPGIRNR

Journal Pre-proof >GhWRKY34 MENFQGFFPTSSSSSTSLSLNLTSSLDYSEFECGKDNDGILGLMADIEVGGAKNENNSLVGTTTESEVKR GKTKKGENKKIRKPRYAFQTRSQVDILDDGYRWRKYGHKAVKNSKFPRSYYRCTHQGCKVKKQVQRLTKD EGIVETTYEGMHSHPIQKTNDNFEHILNQMQIYTSFKSII >GhWRKY36 MEVKEAERIVIAKPVASRPTCSSFKSFSELLAGAINASPPNACSANIVPAIRPKTVRFKPVVNRAPSAMV SSQAEQSRIGISISSDKVSKSDVKPTVVYKPQAKLVSKTTVSLLANMGNFSASNQQRLQSMEAPVEYANW EKMRAQVRPNHHQNAPSQAETDQTSEPSKVGSQNMEEDPKVLPAVANSDRPSYDGYNWRKYGQKQVKGSE YPRSYYKCTHPNCPVKKKVERSLDGQIAEIVYKGEHNHPKPQPPKRNSSQGLGVTSDGTGQDANNSLWSN NHNERNEGSETRAENHSEVRLSVLPAYQVKALAPYEHVTTGGASENSTGLSGECEEASKEGADVESKSKR RKNENQSSEVGTLGECIQEPRVVVQSCTDSEIMGDGFRWRKYGQKVVKGNPYPRSYYRCTNVKCNVRKHV ERASDDPRAFITTYEGKHNHEMPLRNTNHVTAASDPDSNSPAIKDK

-p

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>GhWRKY37 MSSSNEKNDPLYNYDPFQGFNSSTFPFFDYNNLLVYSQAAPTPPFQGLDSSYMNFTDCLHDQNSFSRPFF DMSGSSSDVVGPVHDNTSNSKQTVDDSVRINIDNTENPSTPNSSVSSPSNGRTADEDSSKTNKGSEHEKS KKVNKPKKERQQREPRFAFLTKAEIDVLEDGYRWRKYGQRAVKNSPYPRSYYRCTTQKCGVKKRVERSFQ DPSVVITTYEGRHNHQIPISLRGNNITSSAGFPHEFLAHYFFPSTSNHQEENSIQHQSVALHQHQQQQQL QVSDYGLLHNLVSSSTRKQAP

Jo ur

na

lP

re

>GhWRKY38 MEKEQKDDRFGCEDELLRELLDDESPLFVLQQGTVQAKSKTSGEDSTKRLICAGPKIEDIENALSVKTWN GTQYQVQQQNRINSIQQERGLTKIENKYTLKIKNYGNCMADDGYKWRKYGQKWIKNTPNPRSYYKCRNPR CSAKKQVERSKDEPDTLIITYEGFHLHFAYPYLPLDNHHQPSPDDTISPAKKAKQGISEAESQVPETIAN YTNIEPLPSSIEGCPPEMSLEAGFSQQGLLEDVVPWVIRNPSHNNNISSISSSSSSSNYSRSPPASPYSL STSPFYDPYLDILSINSSIRGYFALAEAGKKLVLNARLSILDSILGNPRIRRLFSSGGPKKRTYENYHPK NKKRIPKANEQKSGGSIGSVESFEEKLEEVQEALGIDPHNYVPVTYVSEVNRFQELLRFGPTLLLLGTFW FMGRRMQSGFEVGDPGGRGGRGIFNIGKAQIKKMDKNAKDKVLFNDVAGCDEAKQEIMEFVHFLKNPKKY EELGAKIPKGAFFVGLSGTGKTLLAKATAGESGVPFLSMSGSDFMKMFVGVGPSRVRSLFQEARQCAPSI IFIDEIDAIGRARGRGGFSGGNGERESTLNQLLVEMRGFGTTSGVVGFDGTNRPNILDRALLRPDIANVF NEAALIAARNGSAVITMEHFEAAVDRVIGGLEKKNKYAMKLCRLQHFWSGFAQYVPNENLLMTKEQLSDV TCMTLGGRDAKQVLLRKISTGAQSDLKKVTKMTYTQVAVYGFSDKVGLLSFPQRDAFEMTKPYSSKTGAI IDSEVRKWVVKAYERTMQLIEEHKEHVTQVAELLLEKVDWIGQKNQIDVAKEAGVKQIVLVGSMGGTNPN HPLNSLGNGNILVWKRKAEQYLADSGVPYTIIRAGGLQDRDGGIRELLVGKDDELLQTETKTIARPDVAE VCIQALKFEEAKFKAFDLASKPEGVGNPTKDFKALFSQITTRF >GhWRKY39 MRFSDENSTAAAESGGGDGGGDKWDSLGFMELLGIHQDFFESFIQQPALILSPPLLQLDTEQKPQAGLQL QPYSTVSEISECLNNPPATPNSSSISSLSNEAANINEQSTNAGDDEEQDKTKKQSYYRCTSSGCGVKKRV ERSFDDTTIIITTYEGKHTHPCPVQPRGSGFGNIGLTPSFMVHHQYQDQQQEEVEQPYMYASLSPLNPCF SPNGSSSASLIRDHGLLQDIVLPIKTRNEANEQV >GhWRKY40 MADEMQEQKPPSKTVASSKRPVITPPPRPFSEALFNGEAGLMGFSPGPMTLVSNFFSDTTDDSKSFSQLL AGAMASPAAQKLTEEQAEGGGVNTGLRFKQNKPAGLAITQQPSSFVMPQGLLSPASLLESPGFSVFSPGP QGPFGMTHQQAVAQVTAQVAQANSHTQIHADTATSLQDSSVMIKQPSDVSQSDQRSQPASDKHASDGYNW RKYGQKQVKGSEFPRSYYKCTHPGCPVKKKVERSLDGQVTEIIYKGQHNHQPPQQNKHAKGNGTSNNRGN SESTSQLHSTNLNIIKEGTCSSMSKKDQESSQATTDLLSGTSDSDDEKDEDEPDTKGRSMEIRVSEPCSS HRTVTEPRIIVQTTSEVDLLDDGYRWRKYGQKVVKGNPYPRSYYKCTTPGCNVRKHVERASTDPKAVITT YEGKHNHDVPAARTSSHNTASSNATQVRTQNIVTDNNHGVNNSVDSTNDRQRPVARLRLKEEQET >GhWRKY41 MAVDLMSFPKMDDQMAVQEAASLGLKGMEHLIRLLSHQSNNVDCTDLADLTVSKFKKVISLLNRTGHARF RRGPIQPLSSSSSPSTSSSASLSVPNSQNNLTLTSAAISTPATINTAVVGTSVISPSSFVQSQPQSLTLD

Journal Pre-proof FTKPNLFGSNGKSTELEFTKESFSVSSNSSFMSSAITGDGSVSNGKQGSSLFLAPALAVSTGKPPLSSTP FKKRCHEHDHSDNVSGKISGSGNSKCHCSKRRKNRVKKVIRVPAISSKIADIPPDEYSWRKYGQKPIKGS PFPRGYYKCSSVRGCPARKHVERAPDDPSMLIVTYEGEHRHTQPAMHENMAPAVGMDWTSSQRDNKQILI >GhWRKY42 MAVELMMSCRNGDSFAAKMEENAVKEAASGLESVEKLIRLLSQTQQQQQTNSNNQEKYQASSISTTSNSL VLEMDCKAEADAAVSRFKRVISLLGRTRTGHARFRRAPVQPDQEQETKVYYATPIQQIPPPVTTAYPLPP PPPPPPHHHHRDFMTVKTGGLERIDSSKTINFSYSSAGNSFLSSLTGDTDSKQPCSSSSAFQITNFSQVS SAGKPPLSSSSSFKRKCSSDNLGSGKCTSGSSGRCHCSKKRKLRSKRVVRVPAISLKMADIPPDDYSWRK YGQKPIKGSPHPRGYYKCSCVRGCPARKHVERALDDPSILVVTYEGEHNHSLSVAETTNLILESS

of

>GhWRKY43 MDAGDSNFNKAGNPFVSEQQVEGNDNVSLETGAESPPPTTTFNDMKLPSPRKGKPIQKRVVSVPIKDVDS SRLKGESAPPSDSWAWRKYGQKPIKGSPYPRGYYRCSSSKGCPARKQVERSRVDPKMLVVTYSYEHNHPW PASRNNTAAAKQAVAAAEAETPTMTTTAAVKPEPSTSQLDTEPESGSEERFADLTDDSILKTRDEFGWLG EMETTSSTVLESQIFSERDNGEGDVGMVFPMREEDELLFADLGELPECSFVFRHQRKMGPQVGIC

re

-p

ro

>GhWRKY44 MWDHEFRNFHCPTDHTCSRVSLSAADPCSYMRFNFTDSLQSPIGYGRSLEKALAFSHSSSSQPPFSSVGT TPNSCISYSSTQDDSDKSNKESQLKELEDGSESCKKGNKLPNKKREKKHKETRFAFMTKSEVDLLEDGYR WRKYGQKAVKNSPYPRSYYRCTTQKCRVKKRVERSFQDPSMVITTYEGQHKHPLPTTVKRSMLFPSFMLR PSPPPPRRRHELFTQMPGDMNNYQAAAGADSLMFAENFSPFRQVPDNGLLQDMVPSAFLKHEPYEQ

lP

>GhWRKY45 MEGERGGVHNYELQVSFSTPQPINEMGFVQFEENQVLSLLALAHSSHHHQISQPLNTTTTTTNSTAMGFT THNNNQVVNTEDPKAANDENNCPSKANDGNNSWWRSSASEKSKLKVRRKLREPRFCFQTRSDVDVLDDGY KWRKYGQKVVKNSLHPRSYYRCTHNNCRVKKRVERLSEDCRMVITTYEGRHNHTPSDDSNSSEHECFTSF

Jo ur

na

>GhWRKY46 MVPSGKCVTDEVGKDKLRSPDAGSHALQHNTDGKIPSSQSDQGGETPLIKSEKDPLSQSEKLGNASSVIT KQTPSLLPGMEGSSPVARERASQDGYNWRKYGQKLVKGNEFVRSYYKCTHPNCRAKKQLERSHDGKMIDT VYVGQHDHPKPLNLPLAVGFAVSVVEERPDKSLQIVVKDKSLHSQMPHQIEPRSGSQPLSSAVSEDVKGA ASKSNRIQNVADSDDDHLVSKRRKKENSNADASPVEKPTNDSRMVIKTFSEVDIVNDGYRWRKYGQKLVK GNPNPRSYYRCSSPGCPVKKHVERASHDAKLVITTYAGHHDHDLPPTRSVTHNTTGVTVHSAAHSDESRT KVEESETVCLDMVVYSGSVAENKSSEQLNGELRTKSDFSGTVCVSLIDAPISGP >GhWRKY47 MEEVEEANKAAIESSHRVLSILSQPQDEVSCRHLMAETGEAVFRFKRVVSLLNSGLGHARVRKLRKLQAP FPQSTLLDTPRCNPSASDYPSKNLHLGCHESSVQELTSHARKSLYLGNPSLELSSNGKHPLQLSQKASLG HYHFLQKQHQMKQQAEMMFRKSNSSGMNLNFDSSSCTPTMSSTRSFLSSLSMDGSVTNLSGGGSAAFHLI GTHRSSDQSSQQKKCYGKAEDGSVKCGTSFICHCSKKRKHRMKRSIKVPAISNKLADIPPDDYSWRKYGQ KPIKGSPHPRGYYKCSSIRGCPARKHVERCLEEPSMLIVTYEGEHNHPRLPSQSTT >GhWRKY48 MEVDWDLHAVVRGCATVTATGNGGVGSGYSMADLHPPQSGVSSFGSEEAFEGHALSSPNQLEELHELYKP FFPKSQPLSPQSTPLSSLSSISFSKEARIEEQQQQSKLSLAASVTTAITAPTNNSATSTSHNSRSKRRKN QLKKVCQVPTEGISSDVWAWRKYGQKPIKGSPYPRGYYRCSTSKGCLARKQVERSRTDPAMFIVTYTAEH NHPAPTHHNSLAGSTRQKPCTSQTPTAGDSVKPFSAKPAGSSSLTTSVDEELMVQNTKAGSREDLVEDEG EDDFGMSDTAVSDDFFEGLEGIADMVTGNCFSDRFQPSCDLPWIASNAATAAGGV >GhWRKY49 MDGQESNPPPPLLSPLNVPFLFPTSLPSSSLHPPLQTQNILPDIDWVSLLSGHENKLMIESGSNNLMSEN GADHHQDAKGANMDKRKGSRIKKTSRPRFAFQTRSEDDILDDGYRWRKYGQKAVKNSIHPRSYYRCTHHT

Journal Pre-proof CNVKKQVQRLSKDTSIVVTTYEGIHNHPCEKLMETLTPLLRQMQFLSSF >GhWRKY50 MEQGALLNELKEGREVTNLLMKHLHLHLPSSHQTGLVLIEKILGSYENALSILNCNGLVVETKPSLSTME SLPSIDHSSPDKRDVFKKRKTSSGWTEQLRVRSGTPLDNPVDDGFCWRKYGQKAILGSNFPRGYYRCTHR HSQGCLAIKQVQKSNDDPTIFEVKYKGRHTCNQTSHFAATYVSVSTESENEGNYSPKRQHQLEVKQKQAQ QAFSSFGSGIKVKTEDLDNREDIFPSFPFPVESEQVKNGSFVEALMENDIMGNLSPAFISPATSESNYFS MSPCHMGSFESDLTDIISAPTSVTNSPIGDLDISSLDKLEFDPSFPFDNPEFFY

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>GhWRKY51 MEKQLRRELRFLHSGDCPTQDSGEPDGALDDSGDHVKPLIKEMDFFPNHNQSHDCHQGRKIDNNNNSPLS SLLDSDVNIGLNLLSSSSGVSRIANEEKPKAEISHVKIALERLNEENRRLRCTLGQITKSYKELQGQLFM AVQKLALGNKTEKKDAVNGTSSPIMSVPQFMDPRPSAALDVNEPSASNDRTQDLSASPGKNMEVVSKEGE HQIPGKHASVEDGSDKTSQSWRPTKCLKLDHSNKEEQVSEVPFRKARVSIRARSEAPLISDGCQWRKYGQ KLAKGNPCPRAYYRCTMAVGCPVRKQVQRCAEDKSILITTYEGNHNHPLPPAATALANTTSAAAAMLLSG STTSKDGLSLPSFGYFPSLPNASTMAPSAAPFPIVTLDLTQGPNAVPFLWPPPSAATFPLPVHGYPQILG HSMLAPPKLSAVSAMQLWQLPVSMVETVTAAIASDPNFSRALAAAISKIIGAPKSNNGGNNNSSNRVATL PGSPQLPQSCTTFSAN

re

-p

>GhWRKY52 MERKEGVKVEDVLGNSSCSVNDFPVQSIFGLSSNEEEKIRSLGFMDLLGVQDLISSPLLEIMAAQQVPSR MATQPPNAFSSTKIGPPHEVFNQPATPNSSTISSASSEAVHDEPAKLDDQEEDQRKTKKQLKPKKTNQKR QREPRFAFMTKSEVDHLEDGYRWRKYGQKAVKNSPFPRSYYRCTTISCNVKKRVERCFSDPSIVVTTYEG QHTHPSPVMPRPNLVGSHLNSAISAASFGMSMQTTPSHYHQHFQQPFTDNLSPLNFGHNGSLNATFLHQK RFCTPGPGPGPSLLKDHGLLQDILPSHMLKEE

na

lP

>GhWRKY53 MIPNSTQLILSHPKEGISFLSLRYVVFSRKVVEKIVVTLKVGENGGKLKSEGPPSDLWSWRKYGQKPIKG SPYPRGYYRCSTSKGCSAKKQVERCKTDASMLIITYTSSHNHPGPDLHCADLKQSPKEPQTTHSDDDKPP TTTLKQLPEVSEKEPEEVKKEQNKTSSTDEDASEEQHFHYLQSPLSFPQNNIISQEDPFNGNLEKTHDTL GFLLDEEPLSCLHITTSSIPKSEENDFFDELEELPISSAFTSLMMRSKFWDEGIPVVPS

Jo ur

>GhWRKY54 MFLSMEKVEEANKAAIESSNRVLSLLSLPKDQLQYSNLMMKTSEAVFKFKKVVSLLNNDLTHSRVRKSKM FRSISPQKIFLESPNCRTILSPKPLQVYPSNLFQKPPLEVKPSQNFSFVHQQQQQQQMQQRLQFQQQQMK YHADMVFGKSNSGINLKFDGSSCTPTMSSAGSFVSSLSLDGSVANLDGNSFHLIGMPHLFGHISQHSRKR CSGIGQDGSMKCGTSGKCHCPKRRKLRIKRSIKVPAITNKVADIPPDEYSWRKYGQKPIKGSPHPRGYYK CSSVRGCPARKHVERCLEDPSMLIVTYEGEHNHSRLLSTHPAHR >GhWRKY56 MEKMGEWEQRSLVSELTQGREVARQLQAHLNGASSLSYDIETREVLVQKIEASYEKALSILNCNVNTSSA SAVVPLTPVIRMPESPNSRCGSPPSEDSDRGFRDSDASKKRKAPRWTQQVRVTPGTALEGTLDDGFNWRK YGQKDILGSRFPRGYYRCTHRNINGCLATKQVQRSDDDPTIFDITYVGTHTCRNVTSHLMSEKNQEQGHS ASSMAAVEPQSSSSSQDLLLNFQKGLKVKTQDLEDQTYASLIPYTSSTSNVVFSWPSVIDNNRNFVSPAT SGTNFDFGTCDDQIIQASAAASVTSSPTVGLDHFHQLDGNFTFDNNGFFRHG >GhWRKY57 MAAGDWDLYAVVRSCTSATNTSTSAVSGNYFSNGNNDSWREDPLACLASLTFEEEDDPLTFPNLKKSGSL QDSYEPLLSDPTTTIISTNRGTDPSSSRSINGVSSGQHHQWQQQFIKPLNTQQQLKKPRKRKNQQKRTVC HVTADNLSSDPWAWRKYGQKPIKGSPYPRNYYRCSSSKGCSARKQVERSNLEPNVFIVTYTGDHTHPRPT HRNSLAGSTRNKLSTFQRAAAPATASCSTSPTTVSVPEGTAADLNNNAGDNGDGDDESADNIAPEMSPEA VSDAEDDLLIPDVHVDDDLFKGLEELVGDAGGGSSSITGNGLGNSSAFGDKFSSWGTGSSASAASGGGW >GhWRKY58

Journal Pre-proof MAEDWDLYAVVRSCTSAANTTRAVENNNNENCSSSFEDPLACLASLTFEEDDDPFPFPNLSQLAETGHLQ DSYKPFLPNADPIAAAAAIDPCSSSSHHGGSSSQLHQHQRLQHQQEQPAATGIGSPLTPTSAPLFTFAGF GNQQQVQPQSHQQQQTRAQPPVQTPRSRKRKNQQKRTVCHVTADKLSSDPWAWRKYGQKPIKGSPYPRNY YRCSSSKGCSARKQVERSNLDSDIFIITYTGEHTHPKPTHRNSLAGSTRNKLSSVQKPTTTKDSAAETIP TTSTVSCSSPRSTTSLSPISPTTTLSAPEDTGAAVHKTGDNGGEEESVYMDVESDEGDDDLLIPDVHVDE GLFKGLEELVGSGSNGVGIGPTFGDNFSSWSTGNSAAAGAAASGGC >GhWRKY59 MEDKEEREQKCLINELLQGRELAMQLQAHLNLPSCNETRELLLQKIQASYDKALSLLNYKAIGKSDTIDP QPQSLPVRDDATLQRKSLARWTQQVRVRPGTATEGNLDDGFSWRKYGQKDILGAKYPRGYYRCTHRNVQG CLATKQVQRSDDDPTIFDVAYRGRHTCSNNNQEQGTSTIEPYQQQQHIGNQTYPLFPNYLSLNRKVENDV GDGNLETGYLSPTSSCPNPMGVMNNGSTSVAIESEFTEIIRAATSAMNAPTVGLDFPFGNAELDPNFTFD DHGFFS

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>GhWRKY60 MEHRGNREQGTLISELIQGMELAKQLRVHLDTASSAESMDLLVQKILSSYEKALLILQFSRPMSMGQPQQ NVGATSGVPESPLSINGSPRSDDLDKDNQDTRDVSKKRKMLPRWTDHVRVTSENLLEGTHDDGYSWRKYG QKDILGAKYPRSYYRCTYRHTQDCWATKQVQKSDEDPTIFEITYRGTHTCVHGNQPVPPPASPEKQEYKL TNLNTNNNQQLQSQAGTLGGRREASQR

lP

re

-p

>GhWRKY61 MTSNFNMEHWDWQAVTGETCSNNDAFISYMENPEFSFGPLSFQHGDEDHLMSFPKVFEPNPNVLDELMEE LYKPFCPELNPYTIPVLEDHAEEPECQKQQPLVVSGTNKDSTKPKRSRKKQQHRLVKHVTGDAGLQSDIW SWRKYGQKPIKGSPYPRSYYRCSSSKGCLARKQVERSCSDPRVFIITYTAEHNHGHPTRRNSLAGSTRNK SLTVAKPKNLAYKNEPHEVAESTVLSPAVIKDESVKQESITMEGGKILTPDVMLSDELVESFEGFGDLFV DQFPDLSHELWFMNESATLTGGLLN

Jo ur

na

>GhWRKY62 MAGIDDNVSLIGDWIPPTQSPRTFFTAMLGDDSGSKQVSKAPLENKSEGLFLGENSDKMDVNQGCEPSNE STFEQKSSTRAGLLERMAARAGFNAPRLNTENIRSTDTSLNPEIRSPYLTIPPGLSPTTLLESPVFVSNS LAQPSPTTGKFSFIPNVNDRSCIPESRDKSEDNYFEDINASSFAFKPIAESNSSFLLGAMSKVTPASLPQ QSFSNIEVSVQSENSQPSRSVDPRKLQQQNSNMLSLQADFLRSSTEKDTGSNNSADQRVFDPVGGSVEHS LPLDEPQDEEGDQRVSGDCMAGGAPSEDGYNWRKYGQKQVKGSEYPRSYYKCTHPTCQVKKKVERSHEGH ITEIIYKGAHNHPKPPPNRRSSIGSSNPLIDMQLELPEQTGLQNGTDGEPVWATAQKGTVAATHDWTHEN IEMTSSATICPENGTAPTPNGTHIESGDAVDASSTFSNDEDEDDRGTHGSVSLGYDGEGDESESKRRKIE AYTTEMSGATRAIREPRVVVQTTSEVDILDDGYRWRKYGQKVVKGNPNPRSYYKCTSAGCTVRKHVERAS HDLKSVITTYEGKHNHDVPAARNSSHVNSATSNTATALTSSSVQTHAHRSEPSQLHNSMVRPTSFGPFTL PGRQQLGPSPGFSFGMNQPGLANLAMAGLGPGQPKMPVLPLHPYMPHQRQGNEMGFRLPKGEAKMEPMSD PGLDLSNNTSVYQQLMSRLPLGPQM >GhWRKY64 MLSSVTKARGPGLAEVQQVEWGAKEIHLADDKHIVSELPRCPNCSSLIALYLQGNYELTAIPPLFFQRMQ LLQILDLSRTSIKSLPKSLRKLFALKKLLLQGCDLFMELSPQVGKLKNLEELHLDETQIMGLPKETGKLL KLQLLKVSFYHLCGKKTLKSDILIHPETISNLSQLAELSIDVNPADKRWDDSVEAVLKEVCNSKTLRTLS LYLPTFQLLDYASLIYPSLSHFRFTVGHHKRRIISRVPHEVEAEFRKWDKCLRFVNGETIPTQIKGVLKY STSFFLDHHTTAMNLSEFGIENMKGLKFCLLAECNKMETLIDGEMHYERNEDDQSESDPGSVQHMLESLE YLSIYYMEDLQCIWRGADRFVCMSKLKFLALHACPQLSKIFSLTLLENFINLEEIILEDCPRVTSLVSHA SVKPIMSDKIFLPSLKRLLVLYLPELVSISNGLLIAPKLESIGCYNCPKLKSISKMELSSKTLKIIKGEC EWWEGMNWNETEWGDGPGYLMHIFSPIDNQKDVMTQMVEGRDPHEATIQNEDQQLGDQKPLEVSTQDHRG QCLDYTEERMMGTDVKEPPSGCVFPSNPLCMTYHAPEQARSFTSGNNRSLEDDECFLVPNIVEVDVDEDE PKAKRWNHTENENKGVIGSASKTTRGNRAANQIRSKVLDDGYRWRKWGQKMVKGNPYPRLYYRCLSTCCL AKKYVERDSQDTSFFVTTYHGLHNHDEWNSRGLSKLHSQPCIDHRANNVKDAAEAAKSICPTKEYGDVFQ ASIQDEGAHPDLAPDDGFCWRKYGQKDILGAKYPRRYYRCANGPSQGCFAKKRVKRSSDDPTIFEITYCG KHTCNVMPPTAPSEYQEQGTRIEPQQQHNQLTEENQKQQSQDLLVLPSTPGQCVEQSFNQKSNSGNDQLT ITHEDNNSTIVCQESSPSPSDSSSMGSQLSAAALSTEQLQAQRFDEPAKQNQLYKKHYPPMLGDEVWRLD RIDKNGIIHKRLASEGINTVQDFLKMWVVNPGELRRILGPIMSERKLDHAINHARTCVMGNKYYVFRGSN

Journal Pre-proof YRILLNPICQLMGAEVNGSIYPTHSLSNIDTVYLEKLVRQAYVNWSSLEEIEGISNEIIGPLTQDIMAQR TGANVINTIPSNLRAMPPSGPWLPELPDHPVLMDNSNVLSSPTTGECAVQYLNQKNNSGNDQQTISQEDN NSTIVCQVPPPSQSNSSAMTSELSATALPTGSDLIQSRAPADAHSWHCYGTKGKKRKHRVKRSIKVPSIG NKLVDIPPDDYSWRKYGQKPIKGSPYSRGYYKCSSMRGCPARKHSERCLEEPSMLIVTYEGKHNHPKLPS RATTSS >GhWRKY65 MIPNSTQLILSHPKEGISFLSLRYVVFSRKVVEKIVVTLKVGENGGKLKSEGPPSDLWSWRKYGQKPIKG SPYPRGYYRCSTSKGCSAKKQVERCKTDASMLIITYTSSHNHPGPDLHCADLKQSPKEPQTTHSDDDKPP TTTLKQLPEVSEKEPEEVKKEQNKTSSTDEDASEEQHFHYLQSPLSFPQNNIISQEDPFNGNLEKTHDTL GFLLDEEPLSCLHITTSSIPKSEENDFFDELEELPISSAFTSLMMRSKFWDEGIPVVPS

ro

of

>GhWRKY66 MEGERGGVHNYELQVSFSTPQPINEMGFVQFEENQVLSLLALAHSSHHHQISQPLNTTTTTTTTNSTAMG FTTQNNNQVVNTEDPKAVNDENNCPSKANDGNNSWWRSSASEKRKLKVRRKLREPRFCFQTRSDVDVLDD GYKWRKYGQKVVKNSLHPRSYYRCTHNNCRVKKRVERLSEDCRMVITTYEGRHNHTPSDDSNSSEHECFT SF

re

-p

>GhWRKY67 MEDKEGGKPGTAFTSESSWTFSGGGPDSVSSSVNYFFDKDSSILSEFGWDLQPDHADDFGRFAEIDRTDS ARPELAGNLSASQSCAAADIVVRCSVSGTALNPGGSADVSTSNPSVSSSSSEDLPEKSTGSGGKPPEIPS KVRKKGQKQIRQPRFAFMTKSEVDHLEDGYRWRKYGQKAVKNSPFPRSYYRCTNSKCTVKKRVERSSEDP AIVITTYEGQHCHHGIAFPRGGLICHEAAFAGQLTPPVSQIYHPGVQSHRQMPPSITQSQPHQAPVDVGE PHPEPEPTSQLPTDEGLLGDIVPPGMRGR

Jo ur

na

lP

>GhWRKY68 MEENLKKRSDDRGGAVVKKEKIADSCADDEDAPAKVGDKRPCCENDDKKPSSPGKKDLSSSNNNKVSLKS DTERTEPEYSMASSSTRKEQDDQLESAKAEMGEVREENQRLKMYLNRIMKDYQKLQMQFYDIVGQDSKES QPAENNDRHHQQQEEEEPELVSLTLGRFSSDSKKDGKNKASSSHGKEEERGNEGLSLGLDYKLEASKSEV DDEALPNPSPVNSTQELKEEETWPPSKVLKTTRTGDDEVLQQNPVKKPRVCVRTRCETPTMNDGCQWRKY GQKIAKGNPCPRAYYRCTVAPSCPVRKQVQRCAEDLSVLITTYEGTHNHPLPMSATAMASTTCAAASMLL SGASSSGSNVASLTSTANLHGLNVYLTDNSKSKFYLPNSSLSAASSHPTITLDLTSTPSSSSSFPFNRFS SAYPTTSRYASTSLSFGSSESNTVSWGNGLLSYGSPSTQPYMKNPLNINGRPQSTMENNSIFPSFMQKNN LNPPQQPLPDTIAAATKAITTDPNFQSALAAALTSIIGTGNNNNGGGGSGESLAQRLKWGEQTFAVNGNG CGSSFFNKPPPSTTSQPGSLIFLPPNSLPFSTPKSASTSPGDTGNHTN >GhWRKY69 MSVDLMRYPKVDDEMTIQEVAAQGLKSMEHLIRLPSHQSNHVECTDVADLTVSKFKKVIKLLNRTGHARF RRGPIHSSSSPSTSSASLTIPDSHKDLTLHPVPIATPVTRNPPTVSDPISFVQSQAPCLTLDFTKPNCFS STTRSTELEFTKDSFSISSNSSFMSSAITGDGSISNGKKASSLFFTLEPAASAEKPPLSSAPYKKRCHEH NHSDNISGSGNGKCHCSKRRKNRVKKVIRVPAISSKIADIPADEYSWRKYGQKPIKGSPYPRGYYKCSAF RGCPARKRVERDSDDPSMLIVTYEWEHRHSEPAMEKNMVPAEGLVFESTR >GhWRKY70 MSPDYDKEKVEVLEAELECLQKENEALGLMFETMSRKYIMLHQAYLRQKSNKRQRLEVPAAAGSKASQVF VKTDPRDQSLIVKDGFQWRKYGQKVTKNNPSPRAYFKCFMAPGCPVKKKVQRCVEDKSIVVATYEGQHNH DVDSTAAGKSMLASCSSAIFSGTRSIPFPPLDNPFPPPITLDLTLSGSDLQNHRNLPSFMHDYSTSNDDG NNKKKKIEDYVASLTKDPNFTLALAAAVARSIKTEHPKPSPP >GhWRKY71 MESTWVDTTLDLNINSSHNTIQVLTGALVEELNRIIAENKKLTEMLTVLCERYSSLQNQYMELVSRNSGS DATAATSKKRKAECEDYVPMIGFSGKAESSFSDEDSCKKPKDCIKAKISRAYVRTNPSDNSLIVRDGYQW RKYGQKVTRDNPSPRAYFKCSFAPSCPVKKKVQRSAEDPSILVATYEGEHNHAQHSTPALSSLSPNSGTS NPRSAPVSSSSSAPTKSSPPTVTLELMKPTGLGNDTKNPTQQVDEPAIQQILVQQMAASLTRDPTFTAAL

Journal Pre-proof AAAISGKVLDHKW >GhWRKY72 MFVSLHMDNYQGEGDLNDIVRASGGGPLSCATGHGSNTVPASSWQQFPSDPLNFPSSAMEDHGLNAFGDP FSAMRDPLLQELDVAAGCSAYFSSPNSTDHIINTTSSATVEADTCNFSGSSTMVGQGKLFEGDMPSPCNI FSRIQISASSSKLPVSPCDSPMMAACVSPRGIKAPAVLPSDMINANTSKSCLIDSTGPLQISSPRNLGIK RRLGKSQVKKVVCIPAPAAANSRSSGEVVPSDLWAWRKYGQKPIKGSPYPRGYYRCSSSKGCSARKQVER SRTDPNMLVITYTSEHNHPWPTQRNALAGSTRSQPSKNNNNNAAASKITSPRSQTQNAVTPKEEDLKESS NDALSPTVAAVNSTASASAKEESEDMEKQLEIDDAEFGEGIPETYRPALMAGDSQQPEDFFAELDEIETD PLNLLFPQGFGGGDQDEQKEKKIMDPFGLFDWPPGNNDASF

ro

of

>GhWRKY73 MDTSSWVDTSLDLSPSSDRPLPLDTPKKEPSGNGNLIVFGKKLSMKEESGALVEELNRVNEENKKLTEML AAMCESYNALQSQLTNLMSKIPVKELSPSKNRKSESSSNNNNNDNYNFGIIGNSESSSTDEDSCKKPKEE IIKAKVSRVYVRTEASDTSLVVKDGYQWRKYGQKVTRDNPSPRAYFKCSFAPTCPVKKKVQRSIDDQSVV IATYEGEHNHLPPSQMEPTSGSGHRGSASLNPSGPTMTPDFGKTKPSGSDVARICSSGLPKMDSPQVRQY LVEQMATSLTKDPKFTAALAAAISGRMFQQSPVE

re

-p

>GhWRKY74 MSDHGFRDFYYHGGAFNNIDHDQQSYGGETDHMYRVSSSSIDPSYMSFTDCLHSTSMDYGSLEKAFGLSP SSSELFSSVEGGNRSMMKQHVGADELGGNTGEVTATLNSSVSSSSSEAGCEEDSDKSKKDGQPKGSDDGG ESSKKGNKAKMKGEKKQREPRFAFVTKSEVDQLEDGYRWRKYGQKAVKDSPYPRSYYRCTTQKCTVKKRV ERSFQDPSTVITTYEGQHNHPLPTTLRGSAAGLFPPSMLTPSPLGRPSFPHELLMQMPNYHHQMNNQAPA GSMFAENFSPFQQYVHQHQVPDYGLLQDMVPSTFLKHEP

Jo ur

na

lP

>GhWRKY76 MEKGWGLTLGNSDPVSSFFHNKTNSTAGPVFRLKQHPDMLQFPVSLAGSREDRPGTSSSSSPASDDNRVA VDEVDFFSDKKHGVVDEKKTGTVTVKTETTSHGRDGPSSDHLDVNTGLHLLTVDGGKSSETEDKREKNAL AELQAEFKRVNAENQKLRNMISHVSNNYTALQMHLVTLMQQHRNREAKSTQEHEIQEVKSEVKKDEVVVP RQFMELVPHSGTAETDVMTHPSSEERTRSGSPPNNVEVASKDSIKGRNEISQFNQEERDGKRIDREESPE SDGWGPNKAQKLNPAKPIEQSTEVTMRKARVSVRARSEAPMISDGCQWRKYGQKMAKGNPCPRAYYRCTM AVGCPVRKQVQRCAEDRTILITTYEGNHNHPLPPAAMAMASTTSAAASMLLSGSMPSTDGMMNPNLLART ILPSSSMATISASAPFPTVTLDLTQSPNPLKFQRPPPQFDGPFSGLPQNLASLSAPQLPQAFGQPPFNQS KFSGLQLSKDMGSSQLGHPQLQQPQLPTLADKVSAATAAITTDPSFTAALAAAITSIIGGAHPNSSSNTS NNDK >GhWRKY77 MELNRDDHLDVEDFLVLPHLETHKEPAATCSSLKAKPSVTLSPVNGSCRDLPSNTLCVDYMLQQNYEGYV VERMEGEDAGKFKVAFRTRSCLEIMDDGYRWRKYGKKKIKNNPNPRHYYRCSREGCKVKKRVEREREDEQ FVITTYEGKHNHENPAAATSLPAHQNPLRYH >GhWRKY78 MTVELKMVYGGDGGGSFARKVVETALKEAANSGIQGVEELITLMSKNQELFNPDVSIKTAAPSGSAESAM DIQAVTDKTVNSFKEVISLLDRPRIGHARFRRAPVLHPKQDTEQASKKIQEPETGSSSFQVTKDQVSAFK PFCSTPSYRLPPLPHSRPQSKRSPPLIAKIGVLERNESPSTINFSSSRTLSAGNSFISSSTGNTDSFQPS GFQFTSPSHVPSSGKPPLFSSLKRKCNSLDGAALKYGSFSRRCHCSKKRKSRLKKVIRVPAISNKTADIP PDEYSWRKYGQKPIKGSPHPRGYYRCSSVKGCPARKHVERALDEPMMLIVTYEADHNHSHNSTDVSPALI LESS >GhWRKY79 MEKHHPSPFFHSDDLIRRRSSASDRTHDDSSDDHVKVKPHIMEMDFFSTDHHQDSKINTNGSSSSFDVNT GLNLVSSRTPSEKKSNTEMSVLKIELERLQEENRRLKSMLDHITRNYNELQGQLFMAVQKQALLAINKQQ VNKEGGVKGMSSHPIMSIQQSMDPRRPSTTLDVNEPSATDDKRTELSASSPANATTSTMDVVSKDGNDDH HRTVQIQGKHVFVEDVIDQTCQQSWGTSPKSSPMVDEPKKEEEQVPEVALKKARVSVRARSEAPMIIDGC QWRKYGQKMAKGNPCPRAYYRCTMAVGCPVRKQVQRCAEDKSILITTYEGNHNHHLPPAATAMAKTTCAA

Journal Pre-proof AAMVLSGSTTSNDGQPNSGYFHSSLPYATMATLSASPPFPTITLDLTQGPNPLPFIRPPPSPPTFPLPLH GYPKLSALPAMQIGQQQQQHTASMVETVTAAIASDPNFTAVLAAAISTFMGSPPPPLPPSNNGNNSTTSQ ALPGSPQLPQSCTTFSTN >GhWRKY80 MERKEGLKIEDYVMGNSINSNSSFCDYTGNVFPLQSVFDSCSEEDNNKFSLGFMELLGVQDFTCPLFDVA QQVPVSTKMGSPEQGFNLPATPNSSSVSSGSSEAVNGEPVKVEDDQEEEDDQQKNKTQLKVKKTNQKRQR EPRFAFMTKSEVDHLEDGYRWRKYGQKAVKNSPFPRSYYRCTSTSCNVKKRVERCFTDPSIVVTTYEGQH THPSPLMPRPNLVGSTINSGISAGAAAAFSVPMQRNTQCHYYYQQQQQQQHQHPFANSLSSLNFGHNGPS IDSSFLHERRLCTSGQSLLKDHGLLQDIIPSHMLKEE

-p

ro

of

>GhWRKY81 MEMDLSLKVDSKEKEEDMEESMDSKMQVEEAEAAEDKEVTSMAAAGEVEDVDAPLELSLQGNNKTHELSV LQLEMNRMKEENKVLRKVVEKTMQDYYDLQMKFAVIQKNDHKQDPPIFLSLNGNGNGNGNSSQDQQPIQR SLNIGSHQKHGSSEGDHNDENNEEELGLSLRLQISSSQREREEEHNKEGSEETPNVASEQNKLQPTCLSA ITSHAVSPPNRKARVSVRARCQTATMNDGCQWRKYGQKIAKGNPCPRAYYRCTVAPGCPVRKQVQRCLED MSILITTYEGTHNHPLPVGATAMASTASAAADSFMFLDSSNPLSNGIPNLSQASLPYQNHPHLMKSHRNV MNFTDPSKGSILDLTNNRHVNHHFPTASSSSSHHPSPAFPWMPSRLSNGTSSRQWKSQEDKSLAENVTAI ASDPKFRVAVAAGLRHHSLPLPLPLPLVTTIGHSWKLIFNPFLQKIYGTLEKKDLLPFMRRSPNRRLEAP IPRPISDYDEEDEGSTLPQRYGVQEGQLVCMEACGVARLGGVATALKVLETLRASETVENLAPFGLNAFR WCIGLTRLGIRKFGLGYFVIGLWKQEANIEDNENLEDVHEEADEMGLLKLASMQTMKQTKLPEPCKLLLK LMLMFCYFQVDVFAT

na

lP

re

>GhWRKY82 MAETESQALPTKRHHHHQNDDSEGQEDDDEEEEEEDGLENEEEEEGASRVSESQGAGAAPPSDDAGWKHS QLDTLPNDDPPDGLQVNSTSESKEGSEFKEKVNIADQEVSPSLAVLSAEAPISLSQQAPTSVTQGISSAP TPTLQEQSLSDQKVNGASVPKANQQKSYNLKMVSVVPIVKTPVSDGYNWRKYGQKQVKSPKGSRSYYKCT FSDCQAKKIECSDDTGHVIEVVNKGMHSHEPPRKNNFTRESKIVSSAARASHNIQPIRIFNDSDPSTSSK ESVPETTVNPERKGLFSCGSDGNGDVQVKEELTSEPETKKRQAFLMKKGNTVCSDSVPKARKKPRFVVHA AGDVGISGDGYRWRKYGQKMVKGNPNPRNYYRCTSAGCPVRKHIETAVDNTNAIIITYKGVHDHDMPVPK KRHGQPIAPLVAAAAPASMNNLQHKKTDGGQSQVTSTQWSVGTEGELTGEALDLGGEKAIESARTLLSIG FEIKPC

Jo ur

>GhWRKY83 MEPAWVDTTLDLNINPCFRTNKAMKREFEGDVVESAPVKYESGVLVEELNRVSAENKKLTKMLTVLCEQY YSLQHQFMEFVNKNPEIETAAAATTSSSKKRKAECEDYGANMIGFSGNTETSSSDDGSPKTPKDCIKPKV SRVQVRTNPSDNSLIVRDGYQWRKYGQKVTRDNPCPRAYFKCSFAPSCPVKKKVQRSAEDPSILVATYEG EHNHHEHHHSPPAEISLSSNNNTPSSNTGSGPVSSAPTKASAPTVTLELLQPGLGGDETERAALQIDAPA IQQILVHQMAASLSRDPNFTAALAAAISGRAV >GhWRKY84 MAVDLMSFPKMDDQMAVQEAASLGLKGMEHLIRLLSHQSNNVDCTDLADLTVSKFKKVISLLNRTGHARF RRGPIQPLSSSSSPSTSSSASLSVPNSQNNLTLTSAAISTPATINTAVVGTSVISPSSFVQSQPQSLTLD FTKPNLFGSNGKSTELEFTKESFSVSSNSSFMSSAITGDGSVSNGKQGSSLFLAPALAVSTGKPPLSSTP FKKRCHEHDHSDNVSGKISGSGNSKCHCSKRRKNRVKKVIRVPAISSKIADIPPDEYSWRKYGQKPIKGS PFPRGYYKCSSVRGCPARKHVERAPDDPSMLIVTYEGEHRHTQPAMHENMAPAVGMDWTSSQRDNKQILI >GhWRKY85 MDAGDSNFNKAGNPFVSEQQVEGNDNVSLETGAESPPPTTTFNDMKLPSPRKGKPIQKRVVSVPIKDVDS SRLKGESAPPSDSWAWRKYGQKPIKGSPYPRGYYRCSSSKGCPARKQVERSRVDPKMLVVTYSYEHNHPW PASRNNTAAAKQAVAAAEAETPTMTTTAAVKPEPSTSQLDTEPESGSEERFADLTDDSILKTRDEFGWLG EMETTSSTVLESQIFSERDNGEGDVGMVFPMREEDELLFADLGELPECSFVFRHQRKMGPQVGIC >GhWRKY86

Journal Pre-proof MWDHEFRNFHCPTDHTCSRVSLSAADPCSYMRFNFTDSLQSPIGYGRSLEKALAFSHSSSSQPPFSSVGT TPNSCISYSSTQDDSDKSNKESQLKELEDGSESCKKGNKLPNKKREKKHKETRFAFMTKSEVDLLEDGYR WRKYGQKAVKNSPYPRSYYRCTTQKCRVKKRVERSFQDPSMVITTYEGQHKHPLPTTVKRSMLFPSFMLR PSPPPPRRRHELFTQMPGDMNNYQAAAGADSLMFAENFSPFRQVPDNGLLQDMVPSAFLKHEPYEQ >GhWRKY87 MGKRCGLALHSLSNLFPDNTSSSSRSSTRLNQNPNMFHFPVSLPCSRDDPHDSGIGTSSDENRLPVDELD FFSAATVNVKKENFHADCARLDVNTGLNLLTANAGSDQSTVDDGVSSDVDDKRVKNEVAQLQVELKRMNA ENQKLKDMVNDVSNNYSALQMHLVNLMQQHQRHEINEVKEQEASLIVPRQLMALRPSGIAETDHEPSHSC SEERTPSGSPPNNVQVTSPKDYLKTKNPVSQLDQESSSFRDGKRIGREESPESENLGRNKLQKMNPSKPI DQATEATMRKARVSVRARSEAPMISDGCQWRKYGQKMAKGNPCPRAYYRCTMAVGCPVRKQVQRCAEDRT ILITTYEGNHNHPLPPAAMAMASTTASAATMLLSGSMHSGDGMMNPNLFPRTMLPCSSMATISASAPFPT VTLDLTHSPNALQFQRPNTQFHVPFPGQPQNSTSTSATQFPQVFGQALYNHNQSKFSGLQLSKDMGSSQL GHPQLQQPQLPTLADKVSAATAAITTDPSFTAALAAAITSIIGGAHPNSSSNTSNNNNNNGNTTNSRQ

-p

ro

of

>GhWRKY88 MDTLYPWPEPETLSSNKKRVIQQLAEGQQCATELQVIVLQNNKPSQQTEELVQKILWSFNQTISMLAEAG HHDEVISQNQATCNDDCKSQDSSESSKRSLSAFAKDKRGCYKRKRFAQTSTVVSDKIEDGHAWRKYGQKN ILNSKHPRSYFRCSHKHDQGCSAIKQVQRMEDDTQMYHITYIGTHTCRGQYSSMATPRIDRLSPISKLES EEQTTTPSDVTDLDSMTMWTDVMMGGVGFEADAVSNMYSCTEITCLDLEPVELENGLQFDDPDFA

na

lP

re

>GhWRKY90 MGKGWGLTLDSDPVTSNFFTNKSNSTTVAGGSFLGLKQHHHHHRDMFQFPVSLAGSRDNDRRGTSASSSP SDDTKIPVDEVDFFSDKKVRVVDDVKKESIETPASDLDVNTGLLLLTAGSDQSTVDDGVSSDMEDKRAKN ELAQLQDELKRMNAENQKLKDMLSHVSNNYSALHMHLVTLMQQQQQPPQRNRETQAIQKSKPEEVTPLQI LDLSRPSAAGAEAAETISHSSSEERTRSGTPPSNVEKRVNREESPESESWGPNCKVQNLSSGKPVDQSTE ATMRKARVSVRARSEAPMITDGCQWRKYGQKMAKGNPCPRAYYRCTMAVGCPVRKQVQRCADDRTILITT YEGNHNHPLPPAAMAMGSTTAAAASMLLSGSMPSADGIMNPNLLARAILPCSSSMATISASAPFPTVTLD LTHSPNPLQFQRPPPPPTQFQVPFPGQPAPVSAPQLPQVFGQPLYNNNQSKFSGLQLSQEMGSSQLGRQQ QQQHQSPPQQPPTLADTVSAATAAITNDPNFTAALAAAISSIISGGTHPNSSNNNNRQ

Jo ur

>GhWRKY91 MDSGGSSSSSFRKVRNPFVTDQELEESDNVSSVTGAESPPPSTTKKGKRSMQKRVVSVPIKDVEGSRLKG EGAPPSDSWAWRKYGQKPIKGSPYPRGYYRCSSSKGCPARKQVERSRVNPTMLVITYSCEHNHAWPASRH NNTSAKQAAAAAAGEASESPTKSSTAVKHEPSTSQPDTEPDSGMEDGFACLTEDSILTTGDEFAWFGEME TTSSTVLESPLFSERDNSEADDTAMIFPMREEDESLFADLEELPECSFVFRHQRNVGPQVGIC >GhWRKY92 MSEHEFREFYYHASTMFHNDHVDDMFRVSSSASVDHHPSSYMSFTDCLHSSVDEYGSLEKAFGLSPSSSE VFSSPIEGLNQTTNQQSSEVMAITPNSSISSSSSEAAYCEEDSEKNNKDKQPKGSDDGSKESPKKGMKGA KKKDERKQKEPRFAFVTKSEVDHLEDGYRWRKYGQKAVKNSPYPRSYYRCTTQKCTVKKRVERSFQDPST VITTYEGQHNHPLPTTLRGNAAALFPPSMFAPATARPSFPHELFMQMAAPHQMNNQQANGVAAASMFGEN FSPYHHQYHHHHRQVSDYGLLQDMVPTRFLKHEP >GhWRKY93 MAVELMMGYDGGNSFVRKIEETALNEATTAGIQGVEELIRLMSKSQQLYNQDASSKTSSPSGSEPAMELQ AITDKTVKSFKKVISLLGQPRTGHARFRRAPLSHLDQEVKHQQQDARQPRQKIPESGDSSLQVAKNQVSA FKPFCSSLGHRLPPLPHNHHQSKSSSLLVPKTGLLERNETASAINFSSSPPLSDGNSFMSSLTGDTDSMH PSLSSGFQFTSPSHIPCSGKPHFSSSLKRKCNSKDDAALKCGSSSSRCHCSKKRKSRVKRVIRVPAISTK MADIPPDDYSWRKYGQKPIKGSPHPRGYYKCSSVRGCPARKHVERAMDDPMMLIVTYEGDHNHRHISDSP APSVLESS >GhWRKY94 MAFNSQVFDNFHGDTSNGMLLSDMKTNKLIQAAQVKEVVENGGFVGSETEIKSGGDRNKKKEKKTRKPRY

Journal Pre-proof AFQTRSQVDILDDGYRWRKYGQKAVKNNKFPRSYYRCTHQGCNVKKQVQRLTKDETLVLTTYEGVHTHPI DNPTDDFHHILSQMQIYTPF >GhWRKY95 MEKKKENLISMANLRFSDENPSAVAAAAAAAAATSFGGSIFDMATGAGACDGDTKWGSSLGFMDLLGIHQ DFIAPSLFDSFQPPPILPPPPPSTSSEPLLHQQLDTKQLQALPSPASIVPESSEVLNNPATPNSSSFSSS SNEAANDHEPTKAEDDEEEEQDQDKTKKQLKPKKKNQKRQREPRFAFMTKSEVDHLDDGYRWRKYGQKAV KNSPFPRSYYRCTSAGCGVKKRVERSSDDPSIVVTTYEGQHKHPYPITPRGSIGINMDPSSSFGRSFVVP QPQYLHQQQQLQPYIYNSSPSLNITSITSSGSSFNASIPDFLQDRRFNTPPTSSSASLLRDHGLLQDIVP TQMRKEAKEQ

-p

ro

of

>GhWRKY96 MAASSSSAFINMDNNVNQSTSTNTTWPSFSDHNKFSTQVPNFKSFAPSPTLISPSSAFSPTDFLDSPVLF STSALFSSPTTTGAFGGAQTVNWKNDSNDHQQGIKPKGFDFSFQVPQTQLTLPSTTATTAFQSCSNTVSM EQPAWSLEKTCGEMVQFKPNMQSNTGSQPAGNYTSQYHQTSQYMNNRKVEDGYNWRKYGQKQVKGSENPR SYYKCTYPNCPTKKKVERSLDGQITEIVYKGSHNHPKPQSTRGNRSSSSHSAEISDQSVGTLGNQLGDSF IMQDETSGSIDDDDFDQASSAISNTNGCDDINENEPDAKRWKGENENEGMVGCGSKTVKEPRIVVQTTSD IDILDDGYRWRKYGQKVVKGNPNPRSYYKCTTIGCPVRKHVERASHDLRAVITTYEGKHNHNVPAARGSG YAMNRPSAVTANMPIRPSAVPSQPNNTMYPNSGVSLGMLQNNNNNNTGSFGFSRLGKSIGSSYMSQPHFS DGAFAKDEPRDDSFLDGFLS

na

lP

re

>GhWRKY97 MEVKEAERIVIAKPVASRPTCSSFKSFSELLAGAINASPPNACSANIVPAIRPKTVRFKPVVNRAPSAMV SSQAEQSRIGISISSDKVSKSDVKPTVVYKPQAKLVSKTTVSLLANMGNFSASNQQRLQSMEAPVEYANW EKMRAQVRPNHHQNAPSQAETDQTSEPSKVGSQNMEEDPKVLPAVANSDRPSYDGYNWRKYGQKQVKGSE YPRSYYKCTHPNCPVKKKVERSLDGQIAEIVYKGEHNHPKPQPPKRNSSQGLGVTSDGTGQDANNSLWSN NHNERNEGSETRAENHSEVRLSVLPAYQVKALAPYEHVTTGGASENSTGLSGECEEASKEGADVESKSKR RKNENQSSEVGTLGECIQEPRVVVQSCTDSEIMGDGFRWRKYGQKVVKGNPYPRSYYRCTNVKCNVRKHV ERASDDPRAFITTYEGKHNHEMPLRNTNHVTAASDPDSNSPAIKDK

Jo ur

>GhWRKY98 MEEGGEKHDWVDDDSQEELVRELLDEESPFFVLSQGTIQPKAITSEEEVSKQLVSNVYSGPRIEDIENAL SSVSNWKDQPQQTTRINSMLERNLSKIESKYTLKVRSCGNGMADDGYKWRKYGQKSIKNSPNPRSYYKCT NPRCNAKKQVEKSRDEADTLIITYEGLHLHFVDPYFPPLSDQPFQHDNNEPTKKPRKAISESDESRAFGL PQTMQIQEATTNNINSAVSLEGCPCPQEVASEGLLEDVVPWIIRNPFHNNIISSNSTSCSSSQTRSPPVS PSSLSWSPNYNPWPVFI >GhWRKY99 MENKYQMFFPISSPASSAAYDPLTTNIMAPTSPHVFNNFHGNSSNGLMDLKTDQKFVKKAAEVKEPVDQN GSFVGSEADYHLKSSAATDKKKEKMIRKPRYAFQTRSEVDILEDGYRWRKYGQKAVKNSKFPRSYYRCTH QGCNVKKQVQRLSKDQSIVVTTYEGIHTHPIHNPTDNFEHILSQMHVYTPLFKPIHL >GhWRKY100 MSSNEKNDPQHNYDPFQGFNRSATFPFFDDNSPCIYNQSAATPPPAFDPSYMTFTDCLNGSNVIDYNSFS RAFDMSCSSSDVVSPMDDNTATDNSKKIPTCGDNNNNISENPSTPNSSVSSSSNEDEDSSKAKKGKQPKG CEDHGDDKSKKVNKATKKEKRQKEPRFAFLTKSEIDHLEDGYRWRKYGQKAVKNSPYPRSYYRCTSPKCG VKKRVERSFEDPSVVITTYEGKHNHHIPATLRGHAAGMFSPPIMSGSMGPTFPHEFVTHFLPSPSNNPAA HANSMHHQTQQQQQQQQQQLQVRDYGLLQDLVSSFTRSQGP >GhWRKY101 MDQQQQQHTTLSLILDGCKLAKELEESIGNLGNQLQPEILSKSCDDIINIFATAKQRLNNNAHHHQDPSL FTHHLLHPPQDSSAHHMQTDPSLQEWLKYGVITQAMDMIQSCRPSISMGGEIQAMDVSDSGKAASSSSSQ RSHRSRKDDEEKCKTRVAAPQMGNTDLPPDDNYTWRKYGQKEILGSKYPRAYYRCTHQKMYNCPAKKQVQ RLDNDFYTFEVTYIGQHTCTMSSTAPSIPQPPPLLHDQMVMTQAMVSQPALPPTSSSTTSSSIVPSGSWL

Journal Pre-proof SMEFSLGSGSGGRGSGGAGSSSGGGPATGSRYGRDVEYSVVADMADVMFNSGSSSSNSMDFIFPCAEDKW EAPGSGDKKH >GhWRKY102 MGTLSPWPETLSNNKKRVIQQLVHGQECATQLQILFHNPSEEGGRLSAEEILVQNILTSFNHTLSALSCI DSSEVSQNQTTCNDDSPWCEDRRSEGCSESSKRPGSKDKRGCYKRKRDAEAWTVVSATMEDGQAWRKYGQ KKILNSIHPRSYFRCTRKYEQGCRAMKQVQRLEDDPQMYEITYIGTHTCTDSFRAPQIIPDSESWESQMG SHFQIPTKHHHLNPAAFPSVKQEAKEAAASDITDMDCMVWKEIMGGGGGFEYSLEHGMGCDYGDCRAFQY EFLKPAEIEHDYHFDGSECLIV

of

>GhWRKY103 MESKEGVLKMEEELGNSFSDYFPLQSIFDWSFEGGENMPLGFMELLGVQDLKPPLFDMGQQQAPSAVVNQ ALNPCSTKNERFNYQPATPNSSSISSASSEAVNDEPIQVDDDQEEDQQKTKKQCVCVFGVCDGGGSRLKP KKTNQKSQREPRFAFMTKSEVDHLEDGYRWRKYGQKAVKNSSFPRSYYRCTTTSCNVKKRVERSFTDPSM VITTYEGQHTHPSPIMPRSSLAGGAGSAPYHYRQQRQPFVNTLPPLNFGHSNGSIKAGNFHHERRFCSPG SSLLKDHGLLQDVIPSHMLKEE

re

-p

ro

>GhWRKY104 MERKEGLKIEDYVMGNSINSNSSFCDYTGNVFPLQSVFDSCSEEDNNKFSLGFMELLGVQDFTCPLFDVA QQVPVSTKMGSPEQGFNLPATPNSSSVSSGSSEAVNGEPVKVEDDQEEEDDQQKNKTQLKVKKTNQKRQR EPRFAFMTKSEVDHLEDGYRWRKYGQKAVKNSPFPRSYYRCTSTSCNVKKRVERCFTDPSIVVTTYEGQH THPSPLMPRPNLVGSTINSGISAGAAAAFSVPMQRNTQCHYYYQQQQQQQHQHPFANSLSSLNFGHNGPS IDSSFLHERRLCTSGQSLLKDHGLLQDIIPSHMLKEE

na

lP

>GhWRKY105 MGESQEQKPSSQPSKTVAVSRPTITPPPRLCTDALFNGGLGMMGFSPGPMTLVSNLFPDSDELKSFSQLL AGAMASPAAGKIPNFSPLTTTEGQGGVTGSGGGGDDTGLRFRQNKPAGLVIAQPPPMFTLATGLSPASLL ESPGFSVFSPIAQGPFGMTHQQALAQVTAQAAQAQCSSTAAASLAPVSAFTADMTTNQQMPISLHSSTVT VNEPSDACQSDHRSQPASLIVDKPADDGYNWRKYGQKQVKGSEFPRSYYKCTSPGCPVKKKVERSIDGQV TEIIYKGQHNHQPPPPNKRAKDTGSLNGNPGNQGNSESASQLQSGSSNILMSKKDQESSQATAEHISGMS DSEEAGENEVGVDEKDEDEPDPKRRSTEIRVSEPASSHRTVTEPRIIVQTTSEVDLLDDGYRWRKYGQKV VKGNPYPRSYYKCTTLGCNVRKHVERAATDPKAVITTYEGKHNHNIPAAKTSSHNTSTVMHHKQECRMQ

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>GhWRKY106 MGTLSAWSENLSTKKKKKVIKQLVEGQDYANQLQILLNNNNYNPSQQQQTEHHLSSTKDQLVDKILSSFN QTLAELTSVNVSSSHNQTGSNDDQLVKSEDCSESRRPRPKGKRGCYKRKRAEQARTVVSDTTQDGHGWRK YGHKDILNSKHPRSYFRCTHKYDQGCRAIKQVQRLEEDGSQMYRTTYIGAHTCKNHSFKAPWIMSDSEFR GTCMAPIERHDDRRSNLAPTTPVVKQETKEEMSGTPDDVTDLDSAMWKDFMEFEYCSEPDVMISNVYSCT GIKFRNFEMDFVNGFEFDGSDECVL >GhWRKY108 MAKKGASSKLSAPTRPTITLPPTPTMDSSFSNGSGLSPGPMTLVSAFFSNQDSTENRSFSQLLAGAMASP GSRLPYNSMDGSFMDVCFKDGGEKRSEPLNFGVPNSALFTVPPGLSPSGLLNSPAFYCLSPQSPFGISHQ QALAQVTAQAAIAQTLNHEETSSQMAPSASGLQSSTMEYAEASQYDSKNQLSVAVDKPAEDGYNWRKYGQ KPIKGCEYSRSYYKYTHLKCPVKKKVERSTDGLITEILYKGAHNHEKPPPNKQGMGSSDGNANSQAPELG SQGLTGNLNKVNAVVPVCSIPGKNHESTEAAELPGSSENEEHCDEEGRDERDNNDEPNPKKRNAATEAAI AFSHKTVTEPKIVVQTRSEVELLDDGYRWRNYGQKVVKGNPYPRSYYKCTNEGCNVRKHVERASMDPRAV ITTYEGKHNHDVPTSRNSVHTQLTTICRSQNHTR >GhWRKY111 MEEVEEANKAAVESCHRVLSLLSQPQDQVHYRNLMAETGEAVFKFKRVVSLLNSSLGHARARKLRKLQTP FPQSIFLDYQRHNPTDYPSKDLQLGYHQELSSHAKSSLYLGTPCLELSSNGKNPLPLAQQPPPPAHYHFL HQRGLQLQQHQAEMLSRKSNSGINFNFDSSSCTPTMSSTRSFISSLSIDGSVANMDSGNAFHLIGAPRSS DQGSQHKKKCSGKGEDGSVKCGSSGKCHCSKKRKHRVKRSIKVPAISTKLADIPPDDYSWRKYGQKPIKG SPHPRGYYKCSSMRGCPARKHVERCLEEPSMLIVTYEGEHNHPKLPSQATT

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>GhWRKY112 MSSSNFNPPDSPESDRADQSNFEFPEDWTLDGWLEDYPETIITGPIQFPFNQADEVNNDSARTSGLLQVP ENETARERRDVRERFAFKTKSEVEILDDGYRWRKYGKKWVKNSPNPRNYYRCSIDGCPVKKRVERDKEDP SYVITTYEGIHNHRSVS >GhWRKY113 MFNSIFLTPPNNPQNSTPICTPTMEITSHTHHLHLDDHFLVDPEQSNCKETSTFISSHGKTSADGSCWEV RGCNSFAYQYIQPNYDDKGSANTADINGDGLRFKIAFRTKSDIEIMDDGYRWRKYGKKKVKNNPNPRNYY QCSSEGCKVKKRVEREREDPRFVITTYVGKHNHETLSPSPLSSAFDISPSSHSTTSLPHTLQILHQP

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>GhWRKY114 MHILLFSFIIEMFNNKFGLPNSPESDCADRTKFELPDAYFDGWLHEGYREAMISGTIENPFNQANEVFDE FACNGSLLIPAPDDNRESETVGEMSELRERYAFKTKSEVEILDDGYRWRKYGKKMVKNSPNPRNYYKCLI EGCPVKKRVERDREDPSYVITTYEGIHNH

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>GhWRKY115 MDFSRESSNPNLNITFYPDNLDPVTEFELSDYLMLDGGVFEEDSSSQSIASSEKGLGGGNEISGATSKTP IIKCKTEAREKKLEQRHRVAFITKSEIEVMDDGYKWRKYGKKPVKNSPNPRNYYKCSSGGCNVKKRIERD RDDHSYVITTYEGSHNHYSPFTVYYNQMPPNAWT

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>GhWRKY116 MQNQGLLEEQQEMPTPMGFFSTFPPNNNITSCPPLGCHPLKAFNIAAPDHHHALPTETLLSSLSSPKHRE HFASDFGGPRLLSLQRSSANFWAWGELNECLGSKKNGVDDHLGVSAMKMKRIKARRKVREPRFCIKTMSE VDVLDDGYKWRKYGQKVVKNTQHPRSYYRCTQDNCRVKKRVERLAEDPGMVITTYEGRHVHSPSHDLDDS QPTDSQLNNFFW

Journal Pre-proof Credit Author Statement

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AK,SA and BC conceptualized the research work. AK, HM, and AP performed all experiments and data analysis. AK, SA and BC wrote the manuscript. HM and AP provided technical support and helped in in silico work. All authors approved the manuscript for submission.