Insights into the bioinformatics and transcriptional analysis of the Elongator complexes (ELPs) Gene Family of wheat: TaELPs contribute to wheat abiotic stress tolerance and leaf senescence

Insights into the bioinformatics and transcriptional analysis of the Elongator complexes (ELPs) Gene Family of wheat: TaELPs contribute to wheat abiotic stress tolerance and leaf senescence | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Insights into the bioinformatics and transcriptional analysis of the Elongator complexes (ELPs) Gene Family of wheat: TaELPs contribute to wheat abiotic stress tolerance and leaf senescence Feng Guo, Md Ashraful Islam, Xiujuan Jin, Lili Sun, Kai Zhao, and 6 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-1521902/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Background: Elongator complexes (ELPs) are the protein complexes that promote transcription through histone acetylation in eukaryotic cells and interact with elongating RNA polymerase II (RNAPII). ELPs role in plant growth and development, signal transduction, and response to biotic and abiotic stresses have been confirmed in model plants. However, the functions of the wheat ELP genes are not well documented. Results: The present study was identified 18 members of the ELPs from the wheat genome by a homology search and further bioinformatics and transcriptions patterns in response to different stress conditions were analyzed to dissect their potential regulatory mechanisms in wheat. Gene duplication analysis showed that 18 pairs of ELP paralogous genes were derived from segmental duplication, which was divided into 6 clades by protein phylogenetic and cluster analysis. The orthologous analysis of wheat TaELPs genes showed that TaELP genes may have evolved from orthologous genes of other plant species or closely related plants. Moreover, a variety of Cis -acting regulatory elements (CAREs) related to growth and development, hormone response, biotic and abiotic stresses were identified in the TaELPs promoter region. Publicly available RNA-seq data analysis indicated that TaELPs gene family members were differentially expressed in wheat seedlings, roots, stems, and leaf panicles, as well as under abiotic stresses. Further, the qRT-PCR analysis showed that the transcription of TaELPs was induced under hormone, salt, and drought stress and during leaf senescence. Conclusions: Overall, TaELP genes might be regulated by hormone signaling pathways and responded to abiotic stress and leaf senescence, which could be investigated further as a potential candidate gene for wheat abiotic stress tolerance and yield improvement. Elongator complexes (ELPs) In silico analysis abiotic stress leaf senescence wheat Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 Figure 10 1. Background RNA polymerase II (RNAPII)-related factors can affect both transcription initiation and elongation phases. Although promoter-regulated transcription initiation requires binding of transcription factors, the multi-subunit Srb/mediator complex-bound RNAPII holoenzyme plays an important role in promoters in Saccharomyces cerevisiae RNAPII can still elongate transcripts in the absence of these conditions.[1, 2] and Otero et al found that the Elongator complexes (ELPs) were the main component of the extended C-terminal repeat domain (CTD)-highly phosphorylated RNA polymerase II (RNAPII) holoenzyme, that affected the elongation of RNAPII transcripts through effects on chromatin under specific conditions[3]. ELP was a protein complex composed of 6 subunits (ELP1-ELP6), of which ELP1-ELP3 subunits formed the core complex and ELP4-ELP6 subunits were auxiliary complexes[4]. In S. cerevisiae , the six genes encoding the ELP subunits are all indispensable for yeast growth and affect sensitivity to high salt, caffeine and 6-azouracil phenotypes, suggesting that six subunits are essential elements for studying elongation factor complex function[4-8]. ELP1 (also known as IKI3, IKBKAP, and IKAP), the largest subunit of Elongator (~150 kDa), is highly conserved in eukaryotes and can be involved in pro-inflammatory cytokine signaling as a regulator of IκB kinases [9]. It contained a conserved WD40 domain, a C-terminal basic region and a phosphorylated segment and was involved in promoting tRNA binding and modification [10, 11]. Elp2 (~90 kDa) was the second-largest subunit compared to the other six subunits, it contains two WD40 domains arranged in tandem and can link ELP1 and ELP3.[5, 10]. ELP3 is considered to be the catalytic subunit of the Elongator complex, which is the enzymatic core protein of ELP and contains an N-terminal iron-sulfur (Fe-S) radical SAM (S-adenosylmethionine) domain and a C-terminal GNAT-type histone acetyltransferase (HAT) domain [10, 12, 13]. In the Elongator complex, Elp1, Elp2 and Elp3 form the core complex, ELP4, ELP5 and ELP6 form the accessory complex and they assemble to form a RecA-type ATPase with a hexameric ring structure, involved in tRNA binding and uridine modification at swing position [8, 10, 14, 15]. Elongator complex (ELP) has been shown to be involved in various cellular gene regulation and biological signal transduction, such as tRNA modification, histone modification, DNA demethylation or methylation, tubulin acetylation and exocytosis, etc. (Figure 1) [16, 17]. In the natural environment, the growth and development of plants are suffering from biotic and abiotic adversities all the time. To resist these stresses, plants have developed various immune defense mechanisms [18]. When plants are subjected to external environmental hazards, cells will be instructed to switch from normal growth and development to stress response. After the stress response was completed, the physiological state was restored to normal growth and development. The transformation of these physiological states was inseparable from the reprogramming of transcriptome, and the strength of plant transformation ability was related to the degree of change in initiating the reprogramming of transcriptome, in which ELP complex played an important regulatory role in this process [19, 20]. Nelissen et al (2005) found that Compared with the control, Arabidopsis elo3 (Atelp3) mutants had lower germination and seed setting rates at the seedling stage, slower germination, delayed flowering, abnormal growth of shoot morphological structures of leaves and flowers, reduced leaf area, apical meristem develops irregularly, and the underground primary roots and hypocotyls grow slowly [21-24]. Zhou, X et al (2009) Four Elongator-related mutants were isolated from Arabidopsis ABA-sensitive mutants, and the observed phenotypes all showed slow leaf and root growth, poor development, and increased ABA sensitivity and anthocyanin accumulation [20]. Besides, Chen Z et al (2006) screened mutants related to drought stress, they found that Arabidopsis AtELP1 mutants were highly sensitive to ABA during growth and development and accelerated stomatal closure to resist drought stress under drought stress, indicating that AtELP1 is in the ABA signal transduction plays an important role [25]. Disruption of the Elongator complex in Arabidopsis caused altered plant physiological signaling and enhanced resistance to oxidative stress induced by CsCl and methyl viologen, suggesting that ELP plays a negative regulatory role in the oxidative stress response [20]. In Arabidopsis pathogen sensing, AtELP2 was found to induce defense gene expression and initiate effector-triggered immunity against pathogen invasion [26]. In the screening of Arabidopsis Atelp2 mutants, mutant Atelp3 was found to respond to pathogens similarly to AtELP2 but did not initiate systemic acquired resistance (SAR) defense mechanisms [27]. It is speculated that when plants encounter pathogen invasion, basal immunity and ETI defense mechanisms are induced by the Elongator complex, but not related to SAR, and the Elongator complex may induce the transcription of defense genes through the effect on chromatin [27]. Heterologous expression of Arabidopsis AtELP3 and AtELP4 in tomatoes can significantly increase the resistance of transgenic tomatoes to Pseudomonas syringae without affecting plant growth and development [28]. In wheat, silencing of TaELP4 reduces resistance to Bacillus cereus and significantly reduces the expression of related defense genes such as TaAGC1, TaCPK7-D, TaPAL5 and inhibits chitinase 2 histone acetylation levels [29]. Thus, the above studies have shown that ELP plays an important role in plant growth and development, biotic stress, and abiotic stress, and the same ELP subunit may participate in different signaling pathways by interacting with multiple downstream targets, mediating plant resistance to multiple stresses. However, the research on plant ELP is mainly concentrated on the model crop Arabidopsis , and the study of other crops ELP is relatively less. Wheat is grown all over the world and provides an indispensable caloric resource for human beings [30]. As the world population grows, wheat agricultural production is projected to need to increase by 38% to meet growing food demand [31]. However, little is known about the ELP gene family in wheat. Compared with the genomes of other crop plants, bread wheat is heterologous sixfold, has the largest genome (16Gb genome size; AABBDD genome), and contains a large number of repeats and transposable elements, which has caused great difficulties in mining high-quality candidate genes and breeding of wheat [32, 33]. With the widespread application of genome sequencing in plants, wheat whole genome sequencing and annotation are also published online, which lays the foundation for us to identify and annotate the ELP gene family in wheat and facilitates the exploration of high-quality candidate genes of the ELP gene family [33, 34]. In this study, we used bioinformatics analysis methods to predict and analyze the number of TaELPs genes based on the whole genome of wheat and gene structure, conserved domains, cis -acting elements, evolutionary relationships, subcellular localization, protein-related functions, hormones, and abiotic stress-induced expression patterns were analyzed. This provides a theoretical basis for further research on the biological functions of members of the wheat TaELPs gene family under abiotic stress. 2. Results 2.1. Identification and annotation of wheat TaELPs family members Arabidopsis and rice ELP proteins were used as reference sequences to search the wheat genome database. After HMM and smart analysis, 18 wheat ELP genes (Table 1 and Table S1) were finally identified and named according to their physical positions on the chromosome. The members of the wheat TaELPs family were further annotated by gene ID, position and open reading frame (ORF) length, and protein physicochemical properties. The ORFs of TaELPs ranged from 753 to 3978 bp and the protein length ranged from 250 to 1325 amino acids. The molecular weights of TaELPs ranged from 27.05 to 147.31 KDa, and according to the predicted isoelectric point (PI) values, the PI ranged from 5.30 to 8.97, of which 6 genes were found to be basic (> 7) and 12 genes were found to be acidic (< 7) [ 35 ]. Table 1 Detailed annotations of the TaELPs in wheat. Gene Name Gene ID Splice PC ORF Chromosome Location Introns Exons Length M.W. PI Instability Aliphatic Index GRAVY SL Prediction Variant Chr Strand Start End Chr Length (aa) (KDa) Index TaELP1-A TraesCS1A02G104700.1 1 II 2511 1A reverse 100,291,939 100,297,147 594,102,056 10 11 836 91.40 6.26 44.07 84.67 -0.093 cytosol TaELP1-B TraesCS1B02G116100.1 1 II 2508 1B reverse 136,647,720 136,652,235 689,851,870 9 10 835 91.36 6.23 42.10 86.05 -0.090 cytosol TaELP1-D TraesCS1D02G096900.1 2 II 2535 1D reverse 83,724,571 83,729,227 495,453,186 9 10 844 92.51 6.25 43.44 83.98 -0.085 nucleus TaELP2-A TraesCS2A02G203700.1 1 III 3978 2A reverse 179,670,464 179,675,987 780,798,557 5 6 1325 147.31 5.60 44.21 90.59 -0.163 cytosol, nucleus, plasma membrane TaELP2-B TraesCS2B02G231000.1 1 III 3975 2B reverse 227,082,327 227,087,872 801,256,715 5 6 1324 147.14 5.52 44.05 90.95 -0.161 cytosol, nucleus, plasma membrane TaELP2-D TraesCS2D02G212000.1 1 III 3978 2D forward 170,443,244 170,448,462 651,852,609 5 6 1325 147.17 5.57 42.54 90.60 -0.158 cytosol, nucleus, plasma membrane TaELP3-A TraesCS2A02G320900.1 1 I 1710 2A forward 550,539,215 550,542,666 780,798,557 8 9 569 63.57 8.88 35.59 85.55 -0.310 cytosol TaELP3-B TraesCS2B02G361800.1 1 I 1710 2B reverse 514,861,001 514,865,480 801,256,715 9 10 569 63.61 8.88 35.55 85.89 -0.307 cytosol TaELP3-D TraesCS2D02G341600.1 1 I 1710 2D forward 436,369,113 436,372,717 651,852,609 8 9 569 63.58 8.88 35.55 85.55 -0.312 cytosol TaELP4-A TraesCS4A02G045700.1 1 V 1155 4A forward 37,776,004 37,782,022 744,588,157 9 10 384 42.72 5.36 51.44 84.32 -0.397 cytosol TaELP4-B TraesCS4B02G259300.1 1 V 1155 4B forward 526,726,516 526,729,679 673,617,499 9 10 384 42.71 5.38 53.49 83.05 -0.413 cytosol TaELP4-D TraesCS4D02G259200.1 1 V 1155 4D reverse 428,719,059 428,727,934 509,857,067 9 10 384 42.71 5.30 52.06 85.34 -0.390 cytosol TaELP5-A TraesCS4A02G105200.1 1 VI 759 4A forward 119,080,643 119,083,199 744,588,157 4 5 252 27.05 5.97 31.37 101.83 0.193 cytosol TaELP5-B TraesCS4B02G198800.1 1 VI 753 4B reverse 427,496,766 427,499,136 673,617,499 5 6 250 27.08 5.98 33.83 99.52 0.159 cytosol TaELP5-D TraesCS4D02G199700.1 1 VI 765 4D reverse 346,452,075 346,454,270 509,857,067 4 5 254 27.24 5.91 35.06 97.56 0.163 cytosol TaELP6-A TraesCS7A02G522900.2 2 IV 1152 7A forward 705,684,956 705,687,644 736,706,236 7 8 383 41.20 8.69 53.26 80.84 -0.252 plastid TaELP6-B TraesCS7B02G439900.1 1 IV 1158 7B forward 705,251,306 705,254,066 750,620,385 7 8 385 41.29 8.97 50.04 78.88 -0.250 plastid TaELP6-D TraesCS7D02G512100.1 2 IV 1155 7D forward 613,867,768 613,870,602 638,686,055 7 8 384 41.18 8.69 47.94 79.35 -0.252 plastid PC, Phylogenetic clade; ORF, Open Reading Frame; No, Number; bp, Base pair; Chr, Chromosome; aa, Amino Acid; M.W., MolecularWeight; Pi, Iso electric point; GRAVY, Grand average of hydropathy, SL,Subcellular Localization. The aliphatic amino acid index and instability index were calculated. The aliphatic amino acid index ranged from 79.35 to 101.83 and the instability index ranged from 31.37 to 53.49. The high aliphatic amino acid index of the protein sequence indicates that it can play a role in a wide temperature range, while the instability index indicates whether the protein is stable or unstable[ 35 ]. Among them, 6 genes were stable (instability index 40)[ 36 ]. The calculated hydropathic index (GRAVY) of TaELPs ranges from − 0.085 to 0.193, indicating that they were hydrophilic and can better interact with water [ 36 ]. Subcellular localization prediction of TaELPs genes showed that most TaELPs family members were localized in the cytoplasm, and 3 genes (TaELP6-A, TaELP6-B, TaELP6-D) were localized in the plastid and (TaELP2-A, TaELP2-B, TaELP2-D) were located in the cytoplasm, nucleus, and plasma membrane respectively. 2.2. Chromosomal distribution and gene duplication of wheat TaELPs genes Eighteen TaELPs genes of wheat were located on 12 wheat chromosomes (Fig. 2 A and Table 1 ). TaELPs genes were evenly distributed in A, B, and D subgenomes, each subgenome contained 6 TaELPs genes (Fig. 2 B). 2A, 2B, 2D, 4A, 4B, and 4D all contained 2 genes, while 1A, 1B, 1D, 7A, 7B, and 7D all contained 1 gene (Fig. 2 C). No TaELPs genes were found on chromosomes 3, 5, and 6, suggesting that TaELPs family genes were unevenly distributed throughout the chromosomal of wheat. Gene duplication analysis showed that there were 18 pairs of ELP paralogous genes in the wheat genome (Fig. 2 D, Table S2), all of which were derived from segmental duplication and located at conserved positions in segmental duplication regions on different chromosomes, indicating that segmental duplication in play an important role in the quantitative expansion of wheat ELP genes [ 37 ]. Two segmental duplications occurred on chromosomes 2A, 2B, 2D, 4A, 4B, and 4D, and one segmental duplication occurred on chromosomes 1A, 1B, 1D, 7A, 7B, and 7D (Fig. 1 D, Table S2). It is further speculated what kind of selection the wheat ELP gene has undergone in the evolutionary process. We calculated the nonsynonymous mutation rate (Ka), synonymous mutation rate (Ks) and the ratio of nonsynonymous mutation rate (Ka) to synonymous mutation rate (Ks) (Ka/Ks) (Table S2). The value of Ka/Ks = 1 denotes that genes experienced a neutral selection; 1 indicates a positive selection [ 38 ]. The Ka/Ks values of the 18 pairs of ELP paralogous genes were all less than 1, suggesting that the TaELPs genes all underwent purification selection after fragment duplication, and the divergence time ranged from 1.83 to 8.53 million years ago (MYA). In conclusion, these results indicate the conserved evolution of TaELPs genes. 2.3. Phylogenetic and cluster analysis of wheat TaELPs To further understand the evolutionary relationship and phylogeny of TaELPs and ELPs in other plant species, we constructed a phylogenetic tree of ELP protein sequences from seven plant species (Fig. 2 , Table S3) by neighbor-joining (NJ). Phylogenetic tree results indicated (Fig. 3 ) that ELP proteins were divided into 6 clades. Among them, clade I was the largest, containing 11 members. Clade II to Clade VI each contained 9 members. Each clade contained both monocotyledonous and dicotyledonous ELP proteins, indicating that the structural features of ELP proteins evolved before the separation of monocotyledonous and dicotyledonous plants. Within each clade, wheat ELP proteins were more distantly related to Arabidopsis thaliana and Solanum lycopersicum ; wheat ELP proteins were closely related to Zea mays, Hordeum vulgare, Brachypodium distachyon , and Oryza sativa , indicating that these species were highly conserved in protein sequences and had similar functions, which can further study the close relationship with wheat. The 18 wheat ELP proteins were evenly divided into 6 clades, each of which contained A, B, and D subgenomes, and the protein sequences were clustered together in a phylogenetic tree. We compared the protein sequence similarity of the A, B, and D subgenomes of the same group, and the results showed that the similarity was more than 95% (Table S4). Studies have shown that the protein sequence similarity and identity of gene duplications exceed 70% and 75%, respectively [ 39 ]. By analyzing the protein sequence and the constructed phylogenetic tree (Fig. 3 , Table S4), it was further confirmed that there is a gene duplication event in the wheat TaELPs family genes. 2.4. Orthologous analysis of wheat TaELPs genes To study the evolutionary relationship of the wheat TaELPs gene family, McscanX software was used to visualize the results of collinearity analysis. We selected the dicotyledonous plants ( Arabidopsis and G. max ), monocotyledonous plants ( O. sativa ) and wheat relatives ( Brachypodium distachyon、Triticum dicoccoides and Aegilops tauschii ) to identify orthologous gene pairs of the wheat ELP genes (Fig. 4 , Table S2). We identified a total of 56 orthologous gene pairs of ELP genes (Table S2). No ELP orthologous gene pairs were observed between Arabidopsis and wheat (At-Ta), and only 3 ELP orthologous gene pairs were found between G. max and wheat (Gm-Ta). Thirteen ELP orthologous gene pairs were found between O.sativa and wheat (Os-Ta). We also found that there are 13, 10 and 17 orthologs of ELPs genes between wheat relatives Brachypodium distachyon、Triticum dicoccoides and Aegilops tauschii (Bd-Ta, Td-Ta and Aet-Ta) and wheat. These results suggest that ELP genes in wheat are distantly related to those in dicotyledonous species and are most closely associated with those in Aegilops tauschii , which might be because Aegilops tauschii are widely considered to be the D-genome ancestor of wheat [ 33 , 40 ]. The Ka/Ks ratio indicates the selection pressure of plant genes and can be used to diagnose the evolutionary form of the sequence [ 41 ]. Thus, we calculated the Ka, Ks, Ka/Ks and T values of all orthologous gene pairs in wheat to further investigate the evolutionary trends of the ELP gene family (Table S2). The results showed that the Ka/Ks ratios of all orthologous genes (Ta-Gm, Ta-Os, Ta-Bd, Ta-Td, and Ta-Aet) were less than 1, suggesting that purification selection plays a dominant role in the evolutionary trend of the ELP gene family. According to the divergence time T value calculated from the Ks value, we found that the divergence time of the orthologous genes (Ta-Gm, Ta-Os, Ta-Bd, Ta-Td, and Ta-Aet) was different, among which the orthologous genes (Ta-Gm) had the longest divergence time and had the shortest divergence time with Aegilops tauschii . In conclusion, TaELPs genes in wheat may have evolved from orthologous genes of other plant species or closely related plants. 2.5. Gene structure and conserved motif analysis of TaELPs genes The phylogenetic tree constructed based on the protein sequences of the members of the wheat TaELPs gene family showed that the members of the TaELPs gene family were divided into three groups (GroupA, GroupB and GroupC), and the results were visualized by combining the gene structure and conserved motifs (Fig. 4 ). The gene structure analysis of wheat TaELPs found that introns ranged from 4–10, and exons ranged from 5–11. A maximum of 11 exons were found in the TaELP1-A , while a minimum of five introns were found in the TaELP5-A and TaELP5-D . In the TaELPs members of GroupA, GroupB and GroupC, the exon-intron numbers of genes were relatively close, and the exon-intron structure of most genes was relatively conservative. For example, in Group A, the number of introns and exons of TaELPs genes were mostly 9 and 10, and the number of untranslated regions (UTRs) was relatively close. To further understand the structural diversity of wheat ELPs , we submitted the protein sequences of 18 TaELPs genes to the MEME5.4.1 online website and predicted 10 conserved motifs (Fig. 5 B). The results showed that the number of conserved motifs ranged from 3 to 9. All TaELPs gene family members contained motif 2, while TaELP4-A , TaELP4-B , and TaELP4-D lacked motif 1. Motif 5 was unique to TaELP3-A , TaELP3-B , and TaELP3-D . The same group of ELP proteins contained similar motifs, and they may also have similarities in gene function. For example, Motif 2, Motif 8, Motif 4, Motif 10 and Motif 7 were included in Group A. The differences in the types and numbers of conserved motifs in wheat ELP proteins reflected the structural diversity of these proteins, indicating that they might have different biological functions. 2.6. Protein conservation domain and 3-D protein structure analysis of TaELPs gene Pfam database was utilized to find the important component domains of TaELPs proteins [ 42 ]. The conserved domains of TaELPs were shown in Fig. 6 . TaELP1-A , TaELP1-B , and TaELP1-D contain four WD40(WD domain, G-beta repeat) protein domains; TaELP4-A , TaELP4-B and TaELP4-D contain 1 Elong_Iki1 (Elongator subunit Iki1) domain; TaELP2-A , TaELP2-B and TaELP2-D contain 1 IKI3 domain; TaELP6-A , TaELP6-B , and TaELP6-D contain 1 PAXNEB domain; TaELP3-A , TaELP3-B and TaELP3-D contain a catalytic domain of S-adenosylmethionine (Radical SAM superfamily) and a histone acetyltransferase (Acetyltransferase (GNAT) family) domain; TaELP5-A , TaELP5-B and TaELP5-D all contain an ELP6 (Elongator complex 6) domain, In addition, an Elong_Iki1 (Elongator subunit Iki1) domain was found in TaELP5-B and TaELP5-D . We used SWISS-MODEL to further identify 3-D models of TaELPs proteins[ 35 , 43 ] and the 3-D structure reveals a few key residues linked to biological processes or intended outcomes, (Figure S1). For TaELP1-A , TaELP1-B and TaELP1-D proteins, 3-D structures were analyzed using the template "6qk7.1.B", a template that describes Elongator complex protein 2. TaELP2-A , TaELP2-B and TaELP2-D proteins, 3-D structures were analyzed using the "6qk7.1.A" template, a template describing the Elongator complex protein 1. TaELP3-A , TaELP3-B , and TaELP3-D Protein, the 3-D structure was analyzed using the "6qk7.1.C" template, a description of Elongator complex protein 3, and identified as containing two ligands (1 x 5AD and 1 x SF4), of which, 5AD (5'-DEOXYADENOSINE)9 residues within 4Å and 4 PLIP interactions, SF4(IRON/SULFUR CLUSTER)8 residues within 4Å and 3 PLIP interactions. TaELP4-A , TaELP4-B and TaELP4-D , 3-D structures were analyzed using the"4a8j.1.B" template, "4a8j.1.B" template was a description ELONGATOR COMPLEX PROTEIN 5; TaELP5-A used "4ejs.1.C" template to analyze 3-D structure, TaELP5-B and TaELP5-D proteins, used "4wia.1.A" to analyze 3-D structure; TaELP6-A , TaELP6-B and TaELP6-D proteins, used The "4a8j.1.A" template analyzes 3-D structures. The Residues in the favored region of the Ramachandran plots generated by all TaELPs ranged from 86.39% to 94.32; the Residues in the outlier region ranged from 0.44–6.27%; Coverage of most TaELPs was above 80%, only TaELP4-A , TaELP4- B and TaELP4-D protein coverage was 58% (Table S5). In addition, we also used SOPMA to calculate the secondary structure elements of the protein sequence (Table S6), the results showed that the TaELPs protein α-helix (Alpha helix) ranged from 13.52–44.46%; β-turn (Beta turn) ranged from 3.39–10.24%; Random coil ranged from 30.31–49.94%; Extended strand ranged from 9.66–32.30%. 2.7. Cis-acting element regulation (CARE) analysis of wheat TaELPs genes A total of 91 different CAREs were identified by analyzing the upstream 2000bp promoter region of the TaELPs gene, mainly investigating abiotic stress and defense-related hormone response elements. All of the identified CAREs were divided into five groups according to their known functions (Fig. 7 B, Table S7). Group I contained 48 environmental stress-related CAREs. Among them, 14 different types of abiotic stress response elements, one cis-element involved in low-temperature response (LTR), one cis-acting element (DRE core) regulating cold and dehydration response gene expression, three anaerobic-induced Essential cis-regulatory elements (ARE, GC-motif, plant_AP-2-like), one MYB binding site (MBS) associated with drought induction, and eight stress response-related response elements (such as MYC, as-1, Unnamed__1, WRE3, etc.) (Fig. 7 A, Table S7/S8). 4 cis-acting elements related to wounding and pathogen response (box S, TC-rich repeats, W box, CCAAT-box), the rest were light-responsive elements of different types, such as 3-AF1 binding site, AE-box, Box II, GT1-motif, chs-CMA1a, etc. Group II contained hormone response-related CAREs. There was a total of 13 different types of CAREs that regulate hormone response, such as cis-acting elements involved in abscisic acid response (ABRE, ABRE2), cis-regulatory elements involved in MeJA response (CGGTA-motif, TGACG-motif), cis-regulatory elements (AuxRR-core, TGA-element) involved in auxin response, and cis-acting elements (TCA-element, TGACG-motif) related to salicylic acid response, etc. Group III contained four core cis-acting elements, among which, CAAT-box and TATA-box appear most frequently in all TaELPs genes, indicating that they play an important role in transcription initiation. TATA-box (including TATA and ATTATA-box) and CAAT-box cis-elements are promoter-associated elements that function at the initiation of transcription [ 35 ]. Group IV was plant growth and development-related CAREs, including cis-elements involved in seed-specific expression (AAGAA-motif, RY-element), cis-acting elements involved in cell cycle regulation (MSA-like), and meristem expression associated cis-regulatory elements (CAT-box) and several other CAREs associated with cell division. Group V was a small number of CAREs with unknown functions, which are also commonly found in the promoter sequences of TaELPs genes, indicating that they may also be involved in the regulatory mechanism of TaELPs genes on the environment (Fig. 7 A, 7 B, Table S7). In conclusion, TaELPs genes may be involved in the regulation of the above environmental stress-related, phytohormone responses, and cell growth and development. These transcription factors CAREs play an important role to induce transcription of TaELPs. 2.8. TaELPs expression pattern prediction analysis To further explore the expression patterns of TaELPs genes in different tissues, developmental stages, and abiotic stresses in wheat, we retrieved all wheat mRNA transcription data from the wheat expression database and visualized TPM values with a heat map (Fig. 8 ). The results showed that TaELP3-A, TaELP3-B, TaELP3-D , and TaELP5-D were expressed at high levels in the tissues of seedlings, roots, stems, leaves, and inflorescences at various stages. In addition, we found that TaELP5-D was up-regulated in flag leaves with the prolongation of post-flowering time, while TaELP3-A , TaELP3-B , and TaELP3-D were gradually down-regulated in flag leaves. Therefore, we speculated that TaELPs genes may be related to senescence; However, TaELP1-D and TaELP6-A had lower expression levels in all different tissues and developmental stages of wheat. TaELP4-A 、 TaELP4-B and TaELP4-D were found to be at higher levels expressed in the root, stem, leaf, and spike tissues. TaELP1-A , TaELP1-B , TaELP2-A , TaELP2-B , TaELP2-D , TaELP5-A , TaELP5-B , TaELP6-B , and TaELP6-D had tissue expression specificity in the stem, milk grain stage、stem 1 cm spike、leaf, seven leaf stage and roots, three-leaf stage. TaELP3-A , TaELP3-B , TaELP3-D , and TaELP5-D had the most obvious up-regulation of gene expression after drought treatment 6h (Figure S2); similarly, the gene expression was up-regulated most obviously after heat treatment 6h; After drought and heat stress treatment, gene expression was slightly down-regulated. TaELP3-A , TaELP3-B , and TaELP3-D had higher expression in low-temperature stress. In salt stress, TaELP3-A , TaELP3- B, and TaELP3-D were all up-regulated to varying degrees and showed a downward trend as a whole; TaELP4-A , TaELP4-B , and TaELP4-D had the highest up-regulated expression after salt stress 48h and showed a trend of upward. TaELP5-D showed very low expression in different treatment times of salt stress; the expression patterns of TaELP2-B and TaELP1-D genes showed an upward trend and the up-regulated expression was most obvious at salt stress 48h; The gene expression of other TaELPs genes were slightly up-regulated or down-regulated in different treatment times of salt stress (Figure S2). 2.9. Expression pattern validation analysis of TaELPs Further understand the potential response mechanism of wheat TaELPs gene family in tolerance to abiotic stress, hormone response, and leaf senescence, we detected the transcription patterns of all TaELPs genes in abiotic stresses (drought, salt, and dark treatment), hormones treatments (IAA, SA, ABA), and during leaf senescence. Under drought treatment, TaELP2 exhibited down-regulated transcriptions at most of the time points. The expression of other TaELPs genes was upregulated to varying degrees at different times of drought treatment. Among them, TaELP3 , TaELP1 , and TaELP4 were significantly upregulated, And TaELP3 significant upregulation was observed after 6h and 72h of drought treatment, The overall trend of TaELP1was upregulated, and significant upregulation was observed from 24h to 48h. The expression of TaELP4 was most significantly upregulated after 12h of drought treatment (Fig. 9 ). Under salt stress, TaELP3 , TaELP1 , and TaELP6 were slightly up-regulated or down-regulated compared with the control (0h); TaELP4 was significantly down-regulated under salt stress 6h, 12h, 24h, and 48h, but after 72h of treatment, The expression was significantly up-regulated immediately; The expression of TaELP2 and TaELP5 was up-regulated, TaELP2 were significantly upregulated from 48h to 72h, And TaELP5 significant upregulation was observed after 6h and 72h of salt treatment (Fig. 9 ).To explore the induced expression patterns of all members of the TaELPs gene family in plant growth and development, we performed dark treatments at different times (Fig. 9 ). The results showed that the expression of all TaELPs genes was up-regulated to varying degrees in the early or late stage of dark treatment. For example, the expression pattern of the TaELP2 gene showed an up-regulated trend; TaELP3 , TaELP1 , TaELP4 , TaELP6 genes showed an up-regulated trend from rising to decline; TaELP5 significant upregulation was observed after 72h of dark treatment (Fig. 9 ). Under IAA treatment, the expression patterns of most TaELPs genes were down-regulated, and only TaELP3 and TaELP5 genes were up-regulated. TaELP3 was most significantly up-regulated after 48h of IAA treatment; TaELP5 was up-regulated immediately after 6h of IAA treatment and the expression level reached a peak and then showed a downward trend (Fig. 10 A). Under SA treatment, it was observed that only TaELP5 exhibited low expression levels at all time treatment stages and other TaELPs genes were up-regulated to varying degrees. For example, TaELP2 was significantly up-regulated in the early and late stages of SA treatment; the gene expression patterns of TaELP4 and TaELP6 were significantly up-regulated after 12h of SA treatment; TaELP3 and TaELP1 significant upregulation was observed after 24h of SA treatment (Fig. 10 B). Under ABA treatment, the relative expression levels of all TaELPs genes were significantly different (Fig. 10 C). TaELP6 showed a significant upregulation at all-time treatment stages compared to the control. TaELP1 , TaELP2 , TaELP4 , and TaELP5 showed an overall upward trend, and the gene expression was up-regulated most significantly after 72h of treatment. Moreover, TaELP3 showed a significant upregulation only at ABA treatment 6h compared to the control (Fig. 10 C). During leaf senescence, all TaELPs genes were up-regulated to varying degrees in the late senescence (Fig. 9 ). The up-regulation trend of TaELP2 and TaELP6 is the same, and the overall showed a trend from decline to rise, TaELP2 was most significantly expressed at 30 days after flowering, followed by TaELP6 at 10 days after flowering; TaELP5 was significantly up-regulated at 10 days and 19 days after flowering; TaELP1 , TaELP3 , and TaELP4 were up-regulated in the same trend. Overall, members of the wheat TaELPs gene family play important regulatory roles in abiotic stresses, hormones, and leaf senescence. 2.10. Prediction of protein-protein interactions of wheat ELPs To study the interaction between wheat TaELPs and other proteins, a network was constructed using the STRING database (Figure S3, Table S9). Based on the predicted results, we observed that TaELP1, TaELP2, TaELP3 and TaELP6 had protein interactions with a Chromatin associated protein KTI12 (Traes_5BL_92F800E16.1, Traes_5BL_D8ECD483D.2 and Traes_5DL_A9A62BF38.1) and a Diphthamide biosynthesis protein 3 (Traes_7BL_69CD9E49D.2, Traes_7DL_EF5C1F9EA.1). In addition, TaELP3 had protein interactions with Traes_2AS_03ED0D137.1, Traes_2BS_E0BE8F2D1.1 and Traes_2DS_E75C5D4AC.1 which encodes WD40 repeat-containing proteins. No protein interacting with TaELP4 and TaELP5 was found. WD repeats proteins are widely present in eukaryotes and are involved in various cellular behavioral and physiological regulations, such as signal transduction, activation of transcriptional activity, cell growth and development, and control of apoptosis. The presence of WD40 domains or repeated WD40 motifs can act as a scaffold for protein-protein or protein-DNA assembly, play an important role in protein interactions and can act as a mediator of transient interactions between other proteins [ 44 ]. Besides, chromatin-associated protein KTI12 was found to interact with the Elongator complex (ELP) in the process of RNA polymerase II promoting transcription elongation [ 45 ]. 4. Discussion Climate warming is a common challenge for global agricultural development and improving agricultural biomass production. According to the predictions of climate change models, global crop yields have been declining as the climate changes, but the decrease in wheat yields is closely related to abiotic stresses [ 46 ]. One of the effective ways to deal with the current bottleneck of wheat production is to tap more wheat stress resistance gene resources and apply them to the breeding of wheat resistant varieties. Elongator complex (ELP) is an indispensable component of gene transcription in eukaryotes, and can also indirectly participate in cell behaviors such as extracellular secretion, telomere gene silencing and DNA damage repair by modifying tRNA to participate in the translation process, which influenced abiotic and biotic stress responses as well as plant growth and development [ 7 ]. 4.1. Evolution and genetic relationship of TaELPs In higher plants, there was a gap between the number of ELP gene family members in closely genetic relationship plants. A total of 6 ELP genes were identified in Arabidopsis , and whereas, 6 members were identified in rice [ 19 , 47 ], which is more closely related to wheat and fewer numbers that were identified in wheat. In the process of studying plant evolution, it is found that the expansion of gene families is closely related to the occurrence of gene duplication, which may originate from segmental, tandem or whole-genome duplication [ 35 , 41 , 48 , 49 ]. Segment duplication (SD) is common in biological evolution. When the repetitive DNA sequence exceeds 1kbp or the identity is higher than 90%, we consider that the gene has SD on the chromosome [ 50 ]. In this study, it was found that the members of the wheat TaELPs gene family contained 18 pairs of ELP paralogous genes, all of which originated from large segment duplication. There were two fragment duplications were observed on 2A, 2B, 2D, 4A, 4B, and 4D and one fragment duplication was observed on 1A, 1B, 1D, 7A, 7B, and 7D (Fig. 2 D, Table S2). These results suggest that fragment duplication may have a dominant role in the evolution and expansion of the wheat TaELPs gene family. The evolutionary selection pressure (Ka/Ks) and divergence time (MYA) of 18 pairs of TaELPs genes were calculated, and the Ka/Ks < 1 of all duplicated gene pairs indicated that TaELPs belonged to purifying selection in long-term evolution. The phylogeny and cluster analysis of wheat TaELPs protein showed that wheat ELP protein was more closely related to the evolution of monocotyledonous plants. In addition, to further understand the genetic relationship of the wheat TaELPs gene family, we investigated the collinearity relationship of ELPs genes in wheat and other species (including monocotyledonous and dicotyledonous plants). No wheat orthologous gene pairs were found in Arabidopsis , only 3 ELPs gene orthologous gene pairs were found in Glycine max, and most wheat orthologous gene pairs were obtained in Oryza sativa , Brachypodium distachyon , Triticum dicoccoides , and Aegilops tauschii . It has been shown that allohexaploid bread wheat (AABBDD) is produced by crossing the allotetraploid Triticum dicoccoides (BBAA) with the diploid Aegilops tauschii (DD) containing the D genome [ 51 ]. Therefore, we speculated that hybridization was part of the reason for more orthologous gene pairs between wheat and Triticum dicoccoides and Aegilops tauschii . The 56 orthologous gene pairs identified in wheat were all derived from whole-genome duplication or fragment duplication within the genome. We also calculated evolutionary selection pressure (Ka/Ks) and divergence time (MYA) between wheat and other species, with Ka/Ks < 1 for all duplicate gene pairs, suggesting that TaELPs belonged to purification selection in long-term evolution. The results of this study suggest that TaELPs genes in wheat may have evolved from orthologous genes in other plant species and are more closely related to monocotyledonous plants. 4.2. Structural diversity of wheat TaELPs The protein conserved domain analysis showed (Fig. 6 ) that each ELP subunit protein in wheat contains its corresponding conserved domain, which further illustrated the evolutionary and functional conservation of eukaryotic ELPs [ 7 ]. Compared with the previously reported domains of Arabidopsis and rice ELPs and wheat TaELPs, we found that there were significant differences in the number of domains between some subunits, such as OsELP2 and AtELP2 contained 5 and 6 WD40 repeat protein domains, respectively, OsELP6 and AtELP6 contained 2 and 1 Elongation complex protein 6 domains respectively[ 10 , 19 ], while wheat contained 1 Elongation complex protein 6 domain and 4 WD40 repeat protein domains, the size of the domains were similar, and the overall difference was not large. It showed that in different plant species, there were slight differences between the structure and number of ELPs , suggesting that there may be changes in function. The WD40 repeat protein domain contained in ELP2 has been widely reported to participate in a variety of biological processes of plant growth and development, and play an important role in protein-protein and protein-DNA interactions. Plant anthocyanin biosynthesis and abiotic stress response are closely related.[ 44 ]. IKI3 was a chromatin-associated domain that interacts with ELP and contains the WD40 repeat protein during RNA polymerase II-promoted transcription elongation [ 3 ]. ELP6 was an accessory subunit of the Elongator complex, was associated with histone acetylation in the nucleus and tRNA modification in the cytoplasm and was able to catalyze the elongation of transcription by RNAII [ 52 ]. PAXNEB has been found in different eukaryotes. It is a component of RNA polymerase II extension protein subunit and HAP subcomplex, which can catalyze intracellular histone acetylation [ 5 , 8 , 15 , 53 ]. Members of the wheat TaELPs gene family had different domains, suggesting that the six subunits had different potential functions in plants. The gene structure and conserved motif analysis of the wheat TaELPs gene showed (Fig. 5 ) that there were quantitative differences in the exon-intron number of the TaELPs gene, which was similar to the results in Arabidopsis and rice[ 19 ]. Among the 10 identified motifs, all members of the wheat TaELPs gene family contained a conserved motif of MOTIF2. These results suggested that the evolutionary pattern of the wheat TaELPs gene family was relatively conserved. Further, subcellular localization prediction showed that most members of the wheat TaELPs gene family were located in the cytoplasm, and a few were located in the nucleus, plastid and plasma membrane. Subcellular localization results of the Elongator complex have been reported in a variety of plants. Nelissen and colleagues (2010) detected GFP-ELO3 fusion protein in the nucleus by the GFP fusion protein method [ 54 ]. Two years later, Tran et al. found ELP3 red fluorescent protein under the detection of confocal laser scanning microscopy through the epidermal cells of faba bean leaves, and the results again showed that the Elongator complex exists in the nucleus and cytoplasm [ 55 ]. In conclusion, most of the above reports have confirmed that the elongator complex (ELP) is localized in the nucleus and cytoplasm, which is also confirmed by our prediction of the subcellular localization of all TaELPs genes in wheat. 4.3 Transcription analysis of TaELP genes reveals its role in wheat growth, development and abiotic stress tolerance Different CAREs distribution in promoter regions may indicate variations in gene regulation and function [ 35 , 56 ]. Through the analysis of 2000 bp cis-acting elements upstream of the promoter of wheat TaELPs gene family members, we can further understand the process of TaELPs gene-regulating physiological changes in wheat. We found that all TaELPs genes contain more than one cis-acting element in response to abiotic stress. The frequency of drought-induced cis-elements MYC, MYB, MBS, etc, and stress defense response cis-elements STRE, as-1, as-1, ARE, etc, and other stress-related CAREs in all TaELPs genes (Fig. 7 B, Table S8) very high [ 56 ]. MYB and MYC transcription factor binding elements were involved in plant responses to drought, high salt, and low temperature, and regulate the expression of related genes under stress[ 57 , 58 ]. In the study of drought resistance of various crops, it was found that MBS elements generally exist in the promoter sequences of drought resistance-related genes and are closely related to the response to drought [ 59 ] indicating that they play an important role in abiotic stress. In addition to the above abiotic stress response elements, they also contained defense and stress response elements and a variety of biological stress-related transcription factor binding elements. We also found hormone-related elements, including ABRE (abscisic acid-responsive), TCA-element (salicylic acid-responsive), CGTCA-motif (MeJA responsive), p-box, and TATC-box (gibberellin responsive element), etc. ABRE response element was related to ABA-related gene expression, which was regulated by ABA-dependent or ABA-independent in abiotic stress [ 60 ]. indicating that TaELPs may be involved in signaling during the wheat stress response. In addition, the frequency of many cells cycle-related CAREs and CAREs with unknown functions was also high, indicating that ELPs genes in wheat have different functions during wheat development. The cis-acting elements of ELP family genes were closely related to their possible physiological processes, Predicting the cis-acting elements of unknown gene families is helpful to quickly predict gene functions, It is an effective way to screen candidate genes and analyze gene functions by reverse genetics [ 19 , 61 ]. The tissue expression patterns of genes are often closely related to their gene functions. The 18 TaELPs genes were expressed to varying degrees in each stage of seedling, root, stem, leaf and panicle, indicating that they may play an important role in the growth and development of wheat plants [ 62 ]. When plants encounter drought stress, the root system will initiate a stress response mechanism to cope with drought by reducing water evaporation and root growth [ 63 , 64 ]. The published transcriptome data show that TaELP1 , TaELP3 , and TaELP4 were expressed at high levels in roots, and their transcription levels were significantly increased under drought stress, suggesting that TaELP1 , TaELP3 , and TaELP4 may affect the root development of plants and respond to drought stress are closely related. qRT-PCR results showed that the expression levels of TaELP1 , TaELP3 , and TaELP4 were significantly increased under drought stress, indicating that the above TaELPs genes play an important role in the regulatory mechanism of plants resisting drought stress (Fig. 9 ). Soil salinization is a limiting factor for today's agricultural development. When studying salt-tolerant plants, it was found that the root system is very sensitive to salt stress and can quickly initiate a stress response mechanism [ 65 , 66 ]. The results of qRT-PCR showed that the expression levels of TaELP2 and TaELP4 were significantly increased during 48–72 h of salt stress, and the expression levels of TaELP5 were significantly up-regulated in the early and late stages of salt stress (Fig. 9 ). Transcriptome analysis showed that TaELP2 , TaELP4 and TaELP5 were highly expressed in root tissue, and their transcription levels were significantly increased in the late stage of salt stress, indicating that TaELP2 , TaELP4 and TaELP5 genes play an important role in the regulation mechanism of plant salt stress resistance. Studies have shown that Elongator plays an important role in plant growth and development. In the phenotypic observation of Arabidopsis elo/Atelp mutants, it was found that the germination and seed setting rates at the seedling stage were lower, the germination was slower, the flowering was delayed, the morphological structure of leaves and flowers was abnormally developed, and the leaf area was reduced, The apical meristem develops irregularly, and the underground taproot and hypocotyl grow slowly [ 21 , 23 , 54 ].This study found that TaELP3 , TaELP4 , TaELP5 and TaELP6 were highly expressed in flag leaves at different time points after flowering (Fig. 9 ), and their transcription levels were significantly increased under dark stress(Fig. 9 ), qRT-PCR results showed that all TaELPs genes were up-regulated to varying degrees in flag leaves (24 to 30 days after flowering), and TaELP6 was the most significantly up-regulated, suggesting that TaELP3, TaELP4 , TaELP5 are closely related to wheat senescence and TaELP6 may play an important role in wheat senescence (Fig. 9 ). Cell proliferation is closely related to Elongator, and cell proliferation is often induced by plant hormones [ 16 , 67 ]. We performed 3 hormone treatments (IAA, SA and ABA) on all TaELPs in wheat. When studying the expression of ELP-related genes in a variety of plants, it was found that a large number of auxin genes have high differential expression, so we speculate that Elongator may induce the expression of auxin-related genes to control plant growth and development [ 54 ]. Further studies found that auxin-related genes were hypoacetylated at histone H3K14, suggesting that the Elongator complex may interact with RNAPII to catalyze the formation and transcription of chromatin, thereby promoting the expression of auxin-related genes [ 24 , 54 ]. Therefore, we treated all TaELPs with IAA at different times and found that only TaELP3 had a higher expression, suggesting that TaELP3 may be an important regulator of wheat growth and development (Fig. 10 A). Salicylic acid (SA) was a signaling molecule that initiates stress response mechanisms when plants encounter pathogens [ 68 ], SA accumulates after pathogen infection and was critical for activating local and systemic acquired resistance [ 69 ]. Elongator inhibits the expression of CAT3 and other related antioxidant genes and can promote the expression of SA-related genes [ 16 , 20 ]. In all TaELPs treated with SA at different times, all TaELPs except TaELP5 were expressed and TaELP2 was significantly expressed, indicating that wheat TaELP2 may play an irreplaceable role in pathogen defense (Fig. 10 B). Under ABA stress treatment, the expression of TaELP3 was not significantly up-regulated, and the remaining TaELPs genes were up-regulated to varying degrees under ABA treatment at different times (Fig. 10 C). We found that TaELP2 , TaELP4 , TaELP5 and TaELP6 were significantly up-regulated in the early and late stages of ABA stress treatment. Plant hormone ABA can cause plant cell behavior regulation and physiological signal transduction, and affect different stages of plant growth and development. When subjected to abiotic stresses such as drought, salt, and high temperature, ABA can conduct signals to activate plant stress response mechanisms [ 20 , 70 – 73 ]. According to related reports, ABA signaling molecules were involved in RNA-related processes such as RNA splicing, RNA structure stabilization, and RNA elongation, indicating that ABA signaling was closely related to RNA metabolism and regulation [ 20 , 25 , 74 – 78 ]. ABA signaling was associated with the histone acetyltransferase Elongator complex [ 25 ]. Therefore, we analyzed the relationship between ABA stress and the three abiotic stresses. The results showed that the expression levels of TaELP2 and TaELP4 were significantly increased at 48-72h induced by salt stress and ABA, and the expression levels of TaELP5 were significantly increased at 6h and 72h. indicating that TaELP2 , TaELP4 and TaELP5 genes play an important role in the regulatory mechanism of plants resisting salt stress and may exist in the salt stress response signaling pathway dependent on ABA regulation. All the above results showed that TaELP2 , TaELP3 , TaELP4 , and TaELP6 may be important regulators of abiotic stress and leaf senescence in wheat, and also play an important role in signal transduction. Therefore, this temporal and spatial expression pattern of TaELP genes indicates that these ELPs might have a function in different tissues and various developmental stages as well as abiotic stress tolerance in wheat. Through predictive analysis of interactions between wheat TaELPs and other proteins (Figure S3, Table S9), We found that TaELPs interacting proteins play roles in plant growth and development, hormonal and abiotic stress responses. The three interacting proteins contained the WD40 domain. The WD-repeat protein can be used as a scaffold for protein-protein assembly, and may interact with TaELPs to play a role in plants, which can regulate plant growth and development, transcriptional regulation, hormone signaling, and initiate plant stress defense mechanisms [ 44 , 79 ]. The WD40 repeat protein had been extensively studied in Arabidopsis , and it can act as a regulator of plant growth and development to regulate plant-specific development [ 79 ]. Chromatin-associated protein KTI12 is an important regulator of the Elongator complex and is involved in the modification of uridine bases in eukaryotic tRNA, and there is a close physical and functional relationship between them [ 45 , 80 ]. Two stress-related genes PtKTI12A and PtKTI12B were identified in Populus trichocarpa under high temperature and drought stress and their expression levels were analyzed in each specific tissue, and it was found that they were differentially expressed [ 81 ]. The results showed that in the stress response mechanism, KTI12 protein can be induced to express, activate plant resistance to stress and participate in tRNA swing uridine modification [ 81 ]. Thus, TaELPs along with their interacting partners might be required to develop wheat stress tolerance. Overall, these findings are useful in elucidating the specific biological activities of TaELP genes in order to generate high-yielding and stress-tolerant wheat cultivars. 5. Conclusions Wheat is a major grain crop and a staple meal all over the world. Therefore, researchers have intended to improve wheat production, quality, and different stress tolerance. The current study was utilized the biological information methods to identify ELP family genes in wheat and carried out a comprehensive and systematic analysis. The gene structure, amino acid motif, and subcellular localization prediction results of the TaELPs gene family showed that the ELP gene in wheat is highly conserved. The homology analysis between wheat and other plant species showed that the TaELPs gene had a close homology relationship with Aegilops tauschii . Cis -acting element analyzed predicts 14 different types of abiotic stress response elements and 13 hormone response-related CAREs. Transcriptome data combined with qRT-PCR results showed that The TaELP genes have a predominant role in the regulatory mechanism of wheat development and stress tolerance. The findings of this work will aid in understanding the role of ELPs in plant developmental processes and various stress situations, as well as their sequential implementation to boost yield and generate stress-tolerant wheat cultivars. 6. Materials And Methods 6.1. Identification of wheat ELPs family gene members We used 12 ELP genes from Arabidopsis and rice to find members of the ELP gene family in the wheat genome [ 19 , 47 ]. The ELP protein sequences of Arabidopsis and rice were retrieved from the Ensemble Plants database. ELP genes were identified from the whole wheat genome by BLASTp against the most recent wheat entire genes from IWGSC (RefSeq v1.0) ( http://plants.ensembl.org/index.html , E-value 100) and through the Blast Compare Two Seqs tool of TBtools [ 82 ]. Finally, after eliminating duplicated sequences, the output of BLASTp and TBtools were selected for domain analysis. SMART ( http://smart.embl-heidelberg.de/ ) or InterPro ( https://www.ebi.ac.uk/interpro ) or NCBI CDD ( https://ncbi.nlm.gov.Structure/cdd/cdd.shtml ) and HMM scan ( https://www.ebi.ac.uk/Tools/hmmer/search/hmmscan ) were utilized to check for the existence of ELP gene family domains in the remaining sequences. The length, molecular weight, isoelectric point (pI), and gross average value (GRAVY) of the wheat ELP proteins were calculated using ProtParam software ( https://web.expasy.org/protparam/ ). 6.2. Sequence alignment and phylogenetic tree construction Full-length protein sequences from several species were aligned using ClustalW in MEGA X [ 83 ], and all sequences after alignment analysis were imported into MEGA X for constructing a phylogenetic tree using the neighbor-joining (NJ) method [ 84 ] with 1000 bootstrap values [ 85 ]. 6.3. Chromosomal localization, gene duplication and collinearity analysis For chromosomal distribution, ELPs genomic positions were obtained from the EnsemblPlants BioMart ( http://plants.ensembl.org/biomart/martview/ ). The ELPs were numbered according to their ascending chromosomal location and were given a “Ta” prefix. TBtools was used to visualize the TaELPs on the wheat chromosomes. McscanX software was used to investigate tandem and segmental duplications within the TaELP gene family and collinearity between ELPs from wheat and several other species [ 86 ]. The TBtools were used to compute the non-synonymous substitution rate (Ka), synonymous substitution rate (Ks), and the Ka/Ks ratio and the divergence time T was estimated by T = Ks/(2×9.1×10^-5)Mya [ 87 ]. 6.4. Subcellular localization and 3-D Structure Modeling Subcellular localization was predicted using WoLF PSORT ( https://wolfpsort.hgc.jp/ ) and CELLO v.2.5( http://cello.life.nctu.edu.tw ). The SWISS-MODEL ( https://swissmodel.expasy.org/interactive#sequence ) was used to create three-dimensional (3D) protein structure of TaELPs. 6.5. Structure, domain and motif analysis of TaELP genes Wheat genome annotation file (GFF3 format), CDS sequence and protein sequence of ELP gene were retrieved from the Ensemble Plants database ( http://plants.ensembl.org/index.html ). The gene-related information of TaELPs was extracted using the function of Gtf/Gff3 Sequences Extract of TBtools [ 87 ]. The Gene Structure Display Server 2.0 ( http://gsds.gao-lab.org/ ) was used to visualize intron, exon, and untranslated regions. The ELP gene domains were retrieved from the Pfam database and Evolview ( https://www.evolgenius.info/evolview-v2/ ) was used to visualize the gene domains. To anticipate the conserved motifs of TaELPs, we employed the motif-based sequence analysis tools MEME version 5.4.1 ( http://meme-suite.org/tools/meme ) with a maximum section of up to 10 motifs and visualized using TBtools [ 87 ]. 6.6. Analysis of cis-acting regulatory elements (CAREs) and protein interaction networks The 2000 bp upstream sequences of 18 TaELPs genes were retrieved from the Ensemble Plants database ( http://plants.ensembl.org/index.html ) and submitted to the online software PlantCARE ( http://bioinformatics.psb.ugent.be/webtools/plantcare/html/ ) to find CAREs. Subsequently, the TBtools Heatmap was used for data visualization [ 87 ]. The STRING online server ( https://string-db.org/cgi ) was used to predict the protein interaction network of TaELPs. 6.7. Tissue specific expression and stress analysis of wheat TaELPs The wheat (Chinese spring) RNA-Seq data was retrieved from the Wheat Exp database ( http://wheat-expression.com/ ), and the TPM (transcripts per million reads) value was used to evaluate the transcript abundance of wheat TaELPs genes. The MeV tool was used to visualize the expression [ 88 ]. 6.8. Plant Materials and treatments Two winter wheat varieties were used in this study, including Yannong 19 (registration no. 2004003) and Jinmai 39 (registration no. ZM0218364). These varieties have been approved by the Ministry of Agriculture and Rural Affairs of the People's Republic of China and are widely cultivated in the traditional farming systems of Shandong and Shanxi provinces. Yannong 19 and Jinmai 39 seeds were maintained and provided by the Yantai Academy of Agricultural Sciences (Shandong, China) and the Institute of Economic Crops of Shanxi Agricultural University (Shanxi Academy of Agricultural Sciences) (Shanxi, China), respectively. Wheat cultivar, Jinmai 39 was selected to study the expression of TaELPs under abiotic stress (drought stress, salt stress, dark stress) and hormone treatments (ABA, IAA, SA). The Seedlings were grown in an artificial climate box (16h light/8h dark) at 22°C. When the wheat plants grew in the 2–3 leaf stage, they were treated with sterile ddH 2 O (control) or 20% PEG-6000 and 250 mM NaCl solutions, respectively. For abscisic acid (ABA), auxin (IAA), salicylic acid (SA) treatments, wheat plants at the same stage were sprayed with 100 mM ABA, IAA, SA, and 0.1% (v/v) ethanol (control). Yannong 19, a delayed senescence wheat cultivar was cultivated in the field and flag leaf samples were taken at 0, 7, 16, 19, 22, 24, 25, and 30 days after anthesis. All leaves collected were immediately frozen in liquid nitrogen and stored in a -80°C freezer for further RNA extraction. Each of the above experiments was set up with 3 sets of repetitions. 6.9. RNA extraction, cDNA first-strand synthesis and real-time PCR analysis The Quick RNA isolation Kit (Tiangen Biochemical Technology Co., Ltd.) was used to extract RNA according to the manufacturer’s instructions and DNase I treatment was used to remove DNA contamination. Synthesis of the first strand of cDNA was carried out according to the instructions of the kit (Baori Doctor Biotechnology (Beijing) Co., Ltd.). To measure the expression of TaELPs , qRT-PCR analysis was performed with specific primers (Table S1). The Elongation factor 1a ( TaEF-1a ) was used as an internal reference gene (GenBank accession no. Q03033) [ 89 ]. The threshold values (CT) were generated using the ABI PRISM 7500 system (Applied Biosystems, Foster City, CA, USA), and the transcription level of TaELPs were assessed using the comparative 2 -ΔΔCT method [ 90 ]. Abbreviations ELP: Elongator complexes; RNAPII: RNA polymerase II; CAREs: Cis -acting regulatory elements; qRT-PCR: Quantitative real-time PCR; CTD: C-terminal repeat domain; Fe-S: iron-sulfur radical; SAM: S-adenosylmethionine; HAT: histone acetyltransferase; SAR: systemic acquired resistance; ORF: open reading frame; PI: predicted isoelectric point; Ka: nonsynonymous mutation rate; Ks: synonymous mutation rate; ABA: abscisic acid; IAA: auxin; SA: salicylic acid. Declarations Acknowledgments Not applicable. Funding This research was sponsored by the State Key Laboratory of Sustainable Dryland Agriculture, Shanxi Agricultural University (No. 202002-2); Shanxi Agricultural University Academic Recovery Special Project (2020xshf02) and Supported by Scientific and Technological Innovation Programs of Higher Education Institutions in Shanxi (2021L124). Author information Affiliations Feng Guo, Md Ashraful Islam, Xiujuan Jin, Lili Sun, Kai Zhao, Juan Lu, Rongyue Yan, Ning Li, Shuguang Wang and Daizhen Sun. State Key Laboratory of Sustainable Dryland Agriculture, College of Agronomy, Shanxi Agricultural University, Taigu 030801, China. Author contributions FG designed the experiment. DZS and MAI provide advice on experimental protocols. XJJ, LLS, KZ, JL and RYY conducted experiments. FG wrote the manuscript. The manuscript was revised by DZS and MAI. The manuscript was grammatically revised and polished by MAI. All authors have read and agree to the published version of the manuscript. Corresponding authors Correspondence to Daizhen Sun. Ethics approval and consent to participate We have obtained the permissions to collect wheat cultivars, Yannong 19 and Jinmai 39 which were acquired from Yantai Academy of Agricultural Sciences (Shandong, China) and the Institute of Economic Crops of Shanxi Agricultural University (Shanxi, China), respectively and identified by Dr. Daizhen Sun. Jinmai 39 was grown in the greenhouse of Shanxi Agricultural University (SXAU). Yannong 19 was grown in the State Key Laboratory of Sustainable Dryland Agriculture, Shanxi Agricultural University. The research conducted in this study required neither the approval of the ethics committee nor any human or animal subjects. The described field studies do not require specific permission. The site is not privately owned or protected in any way, and field research does not involve endangered or protected species. The collection and experimental research of wheat materials involved in this study were approved by Shanxi Agricultural University (SXAU) in China and comply with the relevant guidelines and regulations. Consent for publication Not applicable. Availability of data and materials All needed genome sequences and genome annotation files of wheat were obtained from Ensemblplants database (https://plants.ensembl.org/Triticum_aestivum/Info/Index). The transcriptome sequencing data of different tissues and various abiotic stresses used in this study were retrieved from the Wheat Exp database (http://wheat-expression.com/) and these RNA-sequencing reads of the Wheat Exp database were previously deposited with NCBI under accession codes PRJEB25639, PRJEB23056, PRJNA436817, SRP133837, PRJEB25640, and PRJEB25593. All data generated in this study are available in public and also included in the article and its Additional files. In addition, all databases used in this study are open in public and the links are as follows: EnsemblPlants: https://plants.ensembl.org/Triticum_aestivum/Info/Index ExPasy: http://web.expasy.org/compute_ pi/ STRING: https://cn.string-db.org/cgi/input?sessionId=bXZTbla6vfa4 Wheat Expression Browser: http://www.wheat-expression.com/ Wheat eFP Browser: http://bar.utoronto.ca/efp_wheat/cgi-bin/efpWeb.cgi GSDS: http://gsds.cbi.pku.edu.cn/ MEME: http://meme-suite.org/ plantCARE: http://bioinformatics.psb.ugent.be/webtools/plantcare/html/ SMART: http://smart.embl-heidelberg.de Competing interests The authors declare no conflict of interest. References Bradsher J, Jackson K, Conaway R, Conaway J: RNA polymerase II transcription factor SIII. I. Identification, purification, and properties . The Journal of biological chemistry 1993, 268 (34):25587-25593. Price D, Sluder A, Greenleaf A: Dynamic interaction between a Drosophila transcription factor and RNA polymerase II . 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Supplementary Files Additionalfile1TableS1.xlsx Additionalfile2TableS2.xlsx Additionalfile3TableS3.xlsx Additionalfile4TableS4.xlsx Additionalfile5FigureS1.docx Additionalfile6TableS5.xlsx Additionalfile7TableS6.xlsx Additionalfile8TableS7.xlsx Additionalfile9TableS8.xlsx Additionalfile10FigureS2.docx Additionalfile11FigureS3.docx Additionalfile12TableS9.xlsx Additionalfile13TableS10.xlsx Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-1521902","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":121624300,"identity":"2af92d89-5ed9-4abd-ba59-68a4a33a7314","order_by":0,"name":"Feng Guo","email":"","orcid":"","institution":"State Key Laboratory of Sustainable Dryland Agriculture, College of Agronomy, Shanxi Agricultural University, Taigu 030801","correspondingAuthor":false,"prefix":"","firstName":"Feng","middleName":"","lastName":"Guo","suffix":""},{"id":121624301,"identity":"37e341f0-f2d5-40b5-ba63-c87b97c31744","order_by":1,"name":"Md Ashraful 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12:59:13","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-1521902/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-1521902/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":24208345,"identity":"0fec4411-2e13-4b33-97f1-d326fddc75eb","added_by":"auto","created_at":"2022-07-22 17:04:03","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":324141,"visible":true,"origin":"","legend":"\u003cp\u003e(A) In the nucleus, Elongator interacts with RNA polymerase (RNAPII) and is responsible for histone acetylation, DNA demethylation and methylation at various genetic loci. (B) Elongator regulates protein translation through tRNA modification.\u003c/p\u003e","description":"","filename":"Figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/7f0ddce4b4c761ea75ffd8f6.png"},{"id":24209165,"identity":"a9ad8902-e89a-4f67-81b6-cd6244286870","added_by":"auto","created_at":"2022-07-22 17:09:03","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1248997,"visible":true,"origin":"","legend":"\u003cp\u003e(A) Illustration of the chromosomal distribution of \u003cem\u003eTaELPs\u003c/em\u003e on wheat chromosomes. The gene name is around the chromosome, and the chromosome numbers of three subgenomes are displayed at the top of each chromosome. (B) Distribution of \u003cem\u003eELP\u003c/em\u003e genes in the three subgenomes. (C) Distribution of ELP genes on 21 chromosomes. (D) Genomic localization and replication event analysis of 18 \u003cem\u003eTaELPs\u003c/em\u003e genes. Light grey lines in the background indicate syntenic blocks within the bread wheat genome. Duplicate events are highlighted with black lines.\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/756cf0184f63e2d910de9103.png"},{"id":24210840,"identity":"ec1fb400-7ab6-46ac-bcc2-4cde30e48e65","added_by":"auto","created_at":"2022-07-22 17:19:03","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":672440,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic analysis of \u003cem\u003eTaELP\u003c/em\u003e proteins. The tree was generated using MEGA X by the neighbor-joining (NJ) method with 1000 bootstrap values. All the species and protein ID used for constructing the tree were presented in Table S3.\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/117b95d23bef61f844b76a55.png"},{"id":24211353,"identity":"4eaa17f1-3181-4969-be4d-352363d9012a","added_by":"auto","created_at":"2022-07-22 17:24:03","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":5338049,"visible":true,"origin":"","legend":"\u003cp\u003eThe syntenic relationship between \u003cem\u003eELPs\u003c/em\u003e of wheat and \u003cem\u003eOryza sativa, Arabidopsis, Brachypodium, soybean, Triticum dicoccoides\u003c/em\u003e and \u003cem\u003eAegilops tauschii\u003c/em\u003e. Collinear blocks in wheat and other plant genomes are represented by grey lines in the background, while syntenic \u003cem\u003eTaELP\u003c/em\u003e gene pairs are highlighted with blue lines.\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/affba3a986e23db8b8166a9b.png"},{"id":24210115,"identity":"8c180cb5-5f05-4463-9ee3-a64395b55e0c","added_by":"auto","created_at":"2022-07-22 17:14:03","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":922315,"visible":true,"origin":"","legend":"\u003cp\u003eGene structure and motif analysis of \u003cem\u003eTaELPs\u003c/em\u003e. (A) Introns are represented by black lines, while exons are represented by grey boxes and untranslated regions (UTRs) are represented by blue boxes. (B) Conserved motifs of members of \u003cem\u003eTaELPs\u003c/em\u003e. Ten patterns were identified using the MEME program and presented with boxes of different colors.\u003c/p\u003e","description":"","filename":"Figure5.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/09b4bcf117ce854292ad6179.png"},{"id":24210843,"identity":"feebe02c-0e67-4684-be1f-87fadfa54314","added_by":"auto","created_at":"2022-07-22 17:19:03","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":135726,"visible":true,"origin":"","legend":"\u003cp\u003eThe conserved domain of \u003cem\u003eTaELP\u003c/em\u003e members was identified from Pfam and SMART databases and presented using Evolview.\u003c/p\u003e","description":"","filename":"Figure6.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/cdd889eda5ff799eda9e76ef.png"},{"id":24210126,"identity":"873fb6b9-a7db-4e0c-bbc4-a9a875ed7592","added_by":"auto","created_at":"2022-07-22 17:14:03","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":6159161,"visible":true,"origin":"","legend":"\u003cp\u003eCis-regulatory elements (CAREs) of the \u003cem\u003eTaELP\u003c/em\u003e gene family. CAREs analysis of the 2 kb upstream region was performed using the PlantCARE online server. (A)Red indicates CREs with higher frequencies, while green indicates CREs with zero frequencies. (B)Grouping of CAREs in the \u003cem\u003eTaELPs\u003c/em\u003e gene family.\u003c/p\u003e","description":"","filename":"Figure7.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/4ac9bb750b18cf20c2935a6f.png"},{"id":24208349,"identity":"3a71fd7d-b92a-4d41-bc03-d34babdb1bff","added_by":"auto","created_at":"2022-07-22 17:04:03","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":1954796,"visible":true,"origin":"","legend":"\u003cp\u003eHeatmap representing the expression patterns of \u003cem\u003eTaELPs\u003c/em\u003e genes. (A)In different organs and developmental stages. (B) The expression of the \u003cem\u003eTaELP3\u003c/em\u003e gene in different organs of wheat, predicted by Wheat eFP Browser.\u003c/p\u003e","description":"","filename":"Figure8.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/bb73b561a8d6bdbeceded822.png"},{"id":24210845,"identity":"5f149290-edb6-44cf-9cbf-a1dc48c0a61d","added_by":"auto","created_at":"2022-07-22 17:19:03","extension":"png","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":456197,"visible":true,"origin":"","legend":"\u003cp\u003eRelative transcript profiles of \u003cem\u003eTaELPs\u003c/em\u003e in response to (A) drought stress, (B) salt stress, (C) dark stress and (D) leaf senescence. The relative transcripts of all genes were analyzed using qRT-PCR. The relative transcript levels of \u003cem\u003eTaELPs\u003c/em\u003e were measured using the comparative threshold (2\u003csup\u003e\u003cstrong\u003e-ΔΔCT\u003c/strong\u003e\u003c/sup\u003e) method. Data normalized with the transcripts of wheat elongation factor, TaEF-1a. The 0 h post-treatment and 0 days after anthesis was used as a control and standardized with 1. Values represent the mean ±SD from three independent biological samples.\u003c/p\u003e","description":"","filename":"Figure9.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/2d93c9bed990870c39436f4c.png"},{"id":24208364,"identity":"b70d413b-811f-4fca-84d7-4440f53aedc0","added_by":"auto","created_at":"2022-07-22 17:04:04","extension":"png","order_by":10,"title":"Figure 10","display":"","copyAsset":false,"role":"figure","size":141709,"visible":true,"origin":"","legend":"\u003cp\u003eRelative transcript profiles of \u003cem\u003eTaELPs\u003c/em\u003e in response to (A) Indole-3-acetic acid (IAA), (B) salicylic acids (SA), (C) abscisic acid (ABA) The relative transcripts of all genes were analyzed using qRT-PCR. The relative transcript levels of \u003cem\u003eTaELPs\u003c/em\u003e were measured using the comparative threshold (2\u003csup\u003e\u003cstrong\u003e-ΔΔCT\u003c/strong\u003e\u003c/sup\u003e) method. Data normalized with the transcripts of wheat elongation factor, \u003cem\u003eTaEF-1a\u003c/em\u003e. The 0 h post-treatment (A, B, C) was used as a control and standardized with 1. Red and green colors denote strong and weak transcription of \u003cem\u003eTaELPs\u003c/em\u003e, respectively. The heat map was generated with TBtools and the tree was constructed with the average linkage clustering method.\u003c/p\u003e","description":"","filename":"Figure10.png","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/27df5aa6b80260e6a1e88c9e.png"},{"id":26380256,"identity":"6c6d1fa7-4c82-41a7-8fb5-81d77f8eef66","added_by":"auto","created_at":"2022-09-13 07:14:33","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":5856379,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/55d8b802-bc8f-49ca-b468-e2a6dbd356ac.pdf"},{"id":24208343,"identity":"8068e280-facb-4571-b51c-1a47b9b0dc9b","added_by":"auto","created_at":"2022-07-22 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17:24:03","extension":"docx","order_by":11,"title":"","display":"","copyAsset":false,"role":"supplement","size":982514,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile11FigureS3.docx","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/aa6fe767e3dd3e45a3095db6.docx"},{"id":24210121,"identity":"5b2788dd-5f45-45a5-88cf-e7d103fbd700","added_by":"auto","created_at":"2022-07-22 17:14:03","extension":"xlsx","order_by":12,"title":"","display":"","copyAsset":false,"role":"supplement","size":23406,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile12TableS9.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/f243020f576cb2d086db0d95.xlsx"},{"id":24208362,"identity":"06c4b488-bd96-4f4b-81d7-d7fa60de87b1","added_by":"auto","created_at":"2022-07-22 17:04:03","extension":"xlsx","order_by":13,"title":"","display":"","copyAsset":false,"role":"supplement","size":11670,"visible":true,"origin":"","legend":"","description":"","filename":"Additionalfile13TableS10.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-1521902/v1/7fd884f4e130e75cf090aef7.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Insights into the bioinformatics and transcriptional analysis of the Elongator complexes (ELPs) Gene Family of wheat: TaELPs contribute to wheat abiotic stress tolerance and leaf senescence","fulltext":[{"header":"1. Background","content":"\u003cp\u003eRNA polymerase II (RNAPII)-related factors can affect both transcription initiation and elongation phases. Although promoter-regulated transcription initiation requires binding of transcription factors, the multi-subunit Srb/mediator complex-bound RNAPII holoenzyme plays an important role in promoters in Saccharomyces cerevisiae RNAPII can still elongate transcripts in the absence of these conditions.[1, 2]\u0026nbsp;and Otero et al found that the Elongator complexes (ELPs) were the main component of the extended C-terminal repeat domain (CTD)-highly phosphorylated RNA polymerase II (RNAPII) holoenzyme, that affected the elongation of RNAPII transcripts through effects on chromatin under specific conditions[3]. ELP was a protein complex composed of 6 subunits (ELP1-ELP6), of which ELP1-ELP3 subunits formed the core complex and ELP4-ELP6 subunits were auxiliary complexes[4]. In \u003cem\u003eS. cerevisiae\u003c/em\u003e, the six genes encoding the ELP subunits are all indispensable for yeast growth and affect sensitivity to high salt, caffeine and 6-azouracil phenotypes, suggesting that six subunits are essential elements for studying elongation factor complex function[4-8]. ELP1 (also known as IKI3, IKBKAP, and IKAP), the largest subunit of Elongator (~150 kDa), is highly conserved in eukaryotes and can be involved in pro-inflammatory cytokine signaling as a regulator of I\u0026kappa;B kinases\u0026nbsp;[9]. It contained a conserved WD40 domain, a C-terminal basic region and a phosphorylated segment and was involved in promoting tRNA binding and modification\u0026nbsp;[10, 11]. Elp2 (~90 kDa) was the second-largest subunit compared to the other six subunits, it contains two WD40 domains arranged in tandem and can link ELP1 and ELP3.[5, 10]. ELP3 is considered to be the catalytic subunit of the Elongator complex, which is the enzymatic core protein of ELP and contains an N-terminal iron-sulfur (Fe-S) radical SAM (S-adenosylmethionine) domain and a C-terminal GNAT-type histone acetyltransferase (HAT) domain\u0026nbsp;[10, 12, 13]. In the Elongator complex, Elp1, Elp2 and Elp3 form the core complex, ELP4, ELP5 and ELP6 form the accessory complex and they assemble to form a RecA-type ATPase with a hexameric ring structure, involved in tRNA binding and uridine modification at swing position\u0026nbsp;[8, 10, 14, 15]. Elongator complex (ELP) has been shown to be involved in various cellular gene regulation and biological signal transduction, such as tRNA modification, histone modification, DNA demethylation or methylation, tubulin acetylation and exocytosis, etc. (Figure 1)\u0026nbsp;[16, 17].\u003c/p\u003e\n\u003cp\u003eIn the natural environment, the growth and development of plants are suffering from biotic and abiotic adversities all the time. To resist these stresses, plants have developed various immune defense mechanisms\u0026nbsp;[18]. When plants are subjected to external environmental hazards, cells will be instructed to switch from normal growth and development to stress response. After the stress response was completed, the physiological state was restored to normal growth and development. The transformation of these physiological states was inseparable from the reprogramming of transcriptome, and the strength of plant transformation ability was related to the degree of change in initiating the reprogramming of transcriptome, in which ELP complex played an important regulatory role in this process\u0026nbsp;[19, 20]. Nelissen et al (2005) found that Compared with the control,\u003cem\u003e\u0026nbsp;Arabidopsis elo3 (Atelp3)\u0026nbsp;\u003c/em\u003emutants had lower germination and seed setting rates at the seedling stage, slower germination, delayed flowering, abnormal growth of shoot morphological structures of leaves and flowers, reduced leaf area, apical meristem develops irregularly, and the underground primary roots and hypocotyls grow slowly\u0026nbsp;[21-24]. Zhou, X et al (2009) Four Elongator-related mutants were isolated from Arabidopsis ABA-sensitive mutants, and the observed phenotypes all showed slow leaf and root growth, poor development, and increased ABA sensitivity and anthocyanin accumulation\u0026nbsp;[20]. Besides, Chen Z et al (2006) screened mutants related to drought stress, they found that \u003cem\u003eArabidopsis AtELP1\u003c/em\u003e mutants were highly sensitive to ABA during growth and development and accelerated stomatal closure to resist drought stress under drought stress, indicating that AtELP1 is in the ABA signal transduction plays an important role\u0026nbsp;[25]. Disruption of the Elongator complex in\u003cem\u003e\u0026nbsp;Arabidopsis\u003c/em\u003e caused altered plant physiological signaling and enhanced resistance to oxidative stress induced by CsCl and methyl viologen, suggesting that ELP plays a negative regulatory role in the oxidative stress response\u0026nbsp;[20]. In \u003cem\u003eArabidopsis\u003c/em\u003e pathogen sensing, AtELP2 was found to induce defense gene expression and initiate effector-triggered immunity against pathogen invasion\u0026nbsp;[26]. In the screening of \u003cem\u003eArabidopsis Atelp2\u003c/em\u003e mutants, mutant Atelp3 was found to respond to pathogens similarly to AtELP2 but did not initiate systemic acquired resistance (SAR) defense mechanisms\u0026nbsp;[27]. It is speculated that when plants encounter pathogen invasion, basal immunity and ETI defense mechanisms are induced by the Elongator complex, but not related to SAR, and the Elongator complex may induce the transcription of defense genes through the effect on chromatin\u0026nbsp;[27]. Heterologous expression of \u003cem\u003eArabidopsis\u003c/em\u003e \u003cem\u003eAtELP3\u003c/em\u003e and \u003cem\u003eAtELP4\u0026nbsp;\u003c/em\u003ein tomatoes can significantly increase the resistance of transgenic tomatoes to \u003cem\u003ePseudomonas syringae\u003c/em\u003e without affecting plant growth and development\u0026nbsp;[28]. In wheat, silencing of TaELP4 reduces resistance to Bacillus cereus and significantly reduces the expression of related defense genes such as TaAGC1, TaCPK7-D, TaPAL5 and inhibits chitinase 2 histone acetylation levels\u0026nbsp;[29]. Thus, the above studies have shown that ELP plays an important role in plant growth and development, biotic stress, and abiotic stress, and the same ELP subunit may participate in different signaling pathways by interacting with multiple downstream targets, mediating plant resistance to multiple stresses.\u003c/p\u003e\n\u003cp\u003eHowever, the research on plant ELP is mainly concentrated on the model crop \u003cem\u003eArabidopsis\u003c/em\u003e, and the study of other crops ELP is relatively less. Wheat is grown all over the world and provides an indispensable caloric resource for human beings\u0026nbsp;[30]. As the world population grows, wheat agricultural production is projected to need to increase by 38% to meet growing food demand\u0026nbsp;[31]. However, little is known about the ELP gene family in wheat. Compared with the genomes of other crop plants, bread wheat is heterologous sixfold, has the largest genome (16Gb genome size; AABBDD genome), and contains a large number of repeats and transposable elements, which has caused great difficulties in mining high-quality candidate genes and breeding of wheat\u0026nbsp;[32, 33]. With the widespread application of genome sequencing in plants, wheat whole genome sequencing and annotation are also published online, which lays the foundation for us to identify and annotate the ELP gene family in wheat and facilitates the exploration of high-quality candidate genes of the ELP gene family\u0026nbsp;[33, 34]. In this study, we used bioinformatics analysis methods to predict and analyze the number of \u003cem\u003eTaELPs\u003c/em\u003e genes based on the whole genome of wheat and gene structure, conserved domains, \u003cem\u003ecis\u003c/em\u003e-acting elements, evolutionary relationships, subcellular localization, protein-related functions, hormones, and abiotic stress-induced expression patterns were analyzed. This provides a theoretical basis for further research on the biological functions of members of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family under abiotic stress.\u003c/p\u003e"},{"header":"2. Results","content":"\u003cdiv id=\"Sec2\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Identification and annotation of wheat \u003cem\u003eTaELPs\u003c/em\u003e family members\u003c/h2\u003e \u003cp\u003e \u003cem\u003eArabidopsis\u003c/em\u003e and rice ELP proteins were used as reference sequences to search the wheat genome database. After HMM and smart analysis, 18 wheat \u003cem\u003eELP\u003c/em\u003e genes (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e and Table S1) were finally identified and named according to their physical positions on the chromosome. The members of the wheat \u003cem\u003eTaELPs\u003c/em\u003e family were further annotated by gene ID, position and open reading frame (ORF) length, and protein physicochemical properties. The ORFs of \u003cem\u003eTaELPs\u003c/em\u003e ranged from 753 to 3978 bp and the protein length ranged from 250 to 1325 amino acids. The molecular weights of \u003cem\u003eTaELPs\u003c/em\u003e ranged from 27.05 to 147.31 KDa, and according to the predicted isoelectric point (PI) values, the PI ranged from 5.30 to 8.97, of which 6 genes were found to be basic (\u0026gt;\u0026thinsp;7) and 12 genes were found to be acidic (\u0026lt;\u0026thinsp;7) [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e].\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eDetailed annotations of the TaELPs in wheat.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"19\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c15\" colnum=\"15\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c16\" colnum=\"16\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c17\" colnum=\"17\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c18\" colnum=\"18\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c19\" colnum=\"19\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eGene Name\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eGene ID\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSplice\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003ePC\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eORF\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"5\" nameend=\"c10\" namest=\"c6\"\u003e \u003cp\u003eChromosome Location\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eIntrons\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c12\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eExons\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c13\"\u003e \u003cp\u003eLength\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c14\"\u003e \u003cp\u003eM.W.\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c15\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003ePI\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c16\"\u003e \u003cp\u003eInstability\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c17\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eAliphatic Index\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c18\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eGRAVY\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c19\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSL Prediction\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003eVariant\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003eChr\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003eStrand\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003eStart\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003eEnd\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003eChr Length\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c13\"\u003e \u003cp\u003e\u003cb\u003e(aa)\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c14\"\u003e \u003cp\u003e\u003cb\u003e(KDa)\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c16\"\u003e \u003cp\u003e\u003cb\u003eIndex\u003c/b\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP1-A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS1A02G104700.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2511\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e100,291,939\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e100,297,147\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e594,102,056\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e836\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e91.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e6.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e44.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e84.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.093\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP1-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS1B02G116100.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2508\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e136,647,720\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e136,652,235\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e689,851,870\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e835\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e91.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e6.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e42.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e86.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.090\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP1-D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS1D02G096900.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2535\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e83,724,571\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e83,729,227\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e495,453,186\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e844\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e92.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e6.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e43.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e83.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.085\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003enucleus\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP2-A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS2A02G203700.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3978\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e179,670,464\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e179,675,987\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e780,798,557\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e1325\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e147.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e44.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e90.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.163\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol, nucleus, plasma membrane\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP2-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS2B02G231000.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3975\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e227,082,327\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e227,087,872\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e801,256,715\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e1324\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e147.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e44.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e90.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.161\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol, nucleus, plasma membrane\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP2-D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS2D02G212000.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIII\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3978\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e170,443,244\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e170,448,462\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e651,852,609\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e1325\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e147.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e42.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e90.60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.158\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol, nucleus, plasma membrane\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP3-A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS2A02G320900.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eI\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1710\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e550,539,215\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e550,542,666\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e780,798,557\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e569\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e63.57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e8.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e35.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e85.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.310\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP3-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS2B02G361800.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eI\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1710\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e514,861,001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e514,865,480\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e801,256,715\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e569\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e63.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e8.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e35.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e85.89\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.307\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP3-D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS2D02G341600.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eI\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1710\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e436,369,113\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e436,372,717\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e651,852,609\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e569\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e63.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e8.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e35.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e85.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.312\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP4-A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS4A02G045700.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1155\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e37,776,004\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e37,782,022\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e744,588,157\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e384\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e42.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e51.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e84.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.397\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP4-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS4B02G259300.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1155\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e526,726,516\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e526,729,679\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e673,617,499\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e384\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e42.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e53.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e83.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.413\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP4-D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS4D02G259200.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1155\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e428,719,059\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e428,727,934\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e509,857,067\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e384\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e42.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e52.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e85.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.390\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP5-A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS4A02G105200.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eVI\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e759\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e119,080,643\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e119,083,199\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e744,588,157\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e252\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e27.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e31.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e101.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e0.193\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP5-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS4B02G198800.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eVI\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e753\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e427,496,766\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e427,499,136\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e673,617,499\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e250\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e27.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e33.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e99.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e0.159\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP5-D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS4D02G199700.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eVI\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e765\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003ereverse\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e346,452,075\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e346,454,270\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e509,857,067\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e254\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e27.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e5.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e35.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e97.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e0.163\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003ecytosol\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP6-A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS7A02G522900.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1152\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7A\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e705,684,956\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e705,687,644\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e736,706,236\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e383\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e41.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e8.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e53.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e80.84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.252\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003eplastid\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP6-B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS7B02G439900.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1158\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7B\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e705,251,306\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e705,254,066\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e750,620,385\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e385\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e41.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e8.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e50.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e78.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.250\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003eplastid\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTaELP6-D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTraesCS7D02G512100.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eIV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1155\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eforward\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e613,867,768\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e613,870,602\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e638,686,055\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e384\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e41.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e8.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c16\"\u003e \u003cp\u003e47.94\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c17\"\u003e \u003cp\u003e79.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c18\"\u003e \u003cp\u003e-0.252\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003eplastid\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"19\"\u003ePC, Phylogenetic clade; ORF, Open Reading Frame; No, Number; bp, Base pair; Chr, Chromosome; aa, Amino Acid; M.W., MolecularWeight; Pi, Iso electric point; GRAVY, Grand average of hydropathy, SL,Subcellular Localization.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThe aliphatic amino acid index and instability index were calculated. The aliphatic amino acid index ranged from 79.35 to 101.83 and the instability index ranged from 31.37 to 53.49. The high aliphatic amino acid index of the protein sequence indicates that it can play a role in a wide temperature range, while the instability index indicates whether the protein is stable or unstable[\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]. Among them, 6 genes were stable (instability index\u0026thinsp;\u0026lt;\u0026thinsp;40), and the remaining \u003cem\u003eTaELPs\u003c/em\u003e genes were unstable (instability index\u0026thinsp;\u0026gt;\u0026thinsp;40)[\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e]. The calculated hydropathic index (GRAVY) of \u003cem\u003eTaELPs\u003c/em\u003e ranges from \u0026minus;\u0026thinsp;0.085 to 0.193, indicating that they were hydrophilic and can better interact with water [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e]. Subcellular localization prediction of \u003cem\u003eTaELPs\u003c/em\u003e genes showed that most \u003cem\u003eTaELPs\u003c/em\u003e family members were localized in the cytoplasm, and 3 genes (TaELP6-A, TaELP6-B, TaELP6-D) were localized in the plastid and (TaELP2-A, TaELP2-B, TaELP2-D) were located in the cytoplasm, nucleus, and plasma membrane respectively.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.2. Chromosomal distribution and gene duplication of wheat \u003cem\u003eTaELPs\u003c/em\u003e genes\u003c/h2\u003e \u003cp\u003eEighteen \u003cem\u003eTaELPs\u003c/em\u003e genes of wheat were located on 12 wheat chromosomes (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA and Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). \u003cem\u003eTaELPs\u003c/em\u003e genes were evenly distributed in A, B, and D subgenomes, each subgenome contained 6 \u003cem\u003eTaELPs\u003c/em\u003e genes (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB). 2A, 2B, 2D, 4A, 4B, and 4D all contained 2 genes, while 1A, 1B, 1D, 7A, 7B, and 7D all contained 1 gene (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eC). No \u003cem\u003eTaELPs\u003c/em\u003e genes were found on chromosomes 3, 5, and 6, suggesting that \u003cem\u003eTaELPs\u003c/em\u003e family genes were unevenly distributed throughout the chromosomal of wheat.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eGene duplication analysis showed that there were 18 pairs of ELP paralogous genes in the wheat genome (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eD, Table S2), all of which were derived from segmental duplication and located at conserved positions in segmental duplication regions on different chromosomes, indicating that segmental duplication in play an important role in the quantitative expansion of wheat ELP genes [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e]. Two segmental duplications occurred on chromosomes 2A, 2B, 2D, 4A, 4B, and 4D, and one segmental duplication occurred on chromosomes 1A, 1B, 1D, 7A, 7B, and 7D (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eD, Table S2). It is further speculated what kind of selection the wheat \u003cem\u003eELP\u003c/em\u003e gene has undergone in the evolutionary process. We calculated the nonsynonymous mutation rate (Ka), synonymous mutation rate (Ks) and the ratio of nonsynonymous mutation rate (Ka) to synonymous mutation rate (Ks) (Ka/Ks) (Table S2). The value of Ka/Ks\u0026thinsp;=\u0026thinsp;1 denotes that genes experienced a neutral selection; \u0026lt;1 suggests a purifying or negative selection, and \u0026gt;\u0026thinsp;1 indicates a positive selection [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]. The Ka/Ks values of the 18 pairs of ELP paralogous genes were all less than 1, suggesting that the \u003cem\u003eTaELPs\u003c/em\u003e genes all underwent purification selection after fragment duplication, and the divergence time ranged from 1.83 to 8.53\u0026nbsp;million years ago (MYA). In conclusion, these results indicate the conserved evolution of \u003cem\u003eTaELPs\u003c/em\u003e genes.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.3. Phylogenetic and cluster analysis of wheat \u003cem\u003eTaELPs\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eTo further understand the evolutionary relationship and phylogeny of \u003cem\u003eTaELPs\u003c/em\u003e and ELPs in other plant species, we constructed a phylogenetic tree of ELP protein sequences from seven plant species (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, Table S3) by neighbor-joining (NJ). Phylogenetic tree results indicated (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) that ELP proteins were divided into 6 clades. Among them, clade I was the largest, containing 11 members. Clade II to Clade VI each contained 9 members. Each clade contained both monocotyledonous and dicotyledonous ELP proteins, indicating that the structural features of ELP proteins evolved before the separation of monocotyledonous and dicotyledonous plants. Within each clade, wheat ELP proteins were more distantly related to \u003cem\u003eArabidopsis thaliana\u003c/em\u003e and \u003cem\u003eSolanum lycopersicum\u003c/em\u003e; wheat ELP proteins were closely related to \u003cem\u003eZea mays, Hordeum vulgare, Brachypodium distachyon\u003c/em\u003e, and \u003cem\u003eOryza sativa\u003c/em\u003e, indicating that these species were highly conserved in protein sequences and had similar functions, which can further study the close relationship with wheat.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe 18 wheat ELP proteins were evenly divided into 6 clades, each of which contained A, B, and D subgenomes, and the protein sequences were clustered together in a phylogenetic tree. We compared the protein sequence similarity of the A, B, and D subgenomes of the same group, and the results showed that the similarity was more than 95% (Table S4). Studies have shown that the protein sequence similarity and identity of gene duplications exceed 70% and 75%, respectively [\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]. By analyzing the protein sequence and the constructed phylogenetic tree (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e, Table S4), it was further confirmed that there is a gene duplication event in the wheat TaELPs family genes.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.4. Orthologous analysis of wheat \u003cem\u003eTaELPs\u003c/em\u003e genes\u003c/h2\u003e \u003cp\u003eTo study the evolutionary relationship of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family, McscanX software was used to visualize the results of collinearity analysis. We selected the dicotyledonous plants (\u003cem\u003eArabidopsis\u003c/em\u003e and \u003cem\u003eG. max\u003c/em\u003e), monocotyledonous plants (\u003cem\u003eO. sativa\u003c/em\u003e) and wheat relatives (\u003cem\u003eBrachypodium distachyon、Triticum dicoccoides\u003c/em\u003e and \u003cem\u003eAegilops tauschii\u003c/em\u003e) to identify orthologous gene pairs of the wheat \u003cem\u003eELP\u003c/em\u003e genes (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e, Table S2). We identified a total of 56 orthologous gene pairs of \u003cem\u003eELP\u003c/em\u003e genes (Table S2). No \u003cem\u003eELP\u003c/em\u003e orthologous gene pairs were observed between \u003cem\u003eArabidopsis\u003c/em\u003e and wheat (At-Ta), and only 3 \u003cem\u003eELP\u003c/em\u003e orthologous gene pairs were found between \u003cem\u003eG. max\u003c/em\u003e and wheat (Gm-Ta). Thirteen \u003cem\u003eELP\u003c/em\u003e orthologous gene pairs were found between \u003cem\u003eO.sativa\u003c/em\u003e and wheat (Os-Ta). We also found that there are 13, 10 and 17 orthologs of ELPs genes between wheat relatives \u003cem\u003eBrachypodium distachyon、Triticum dicoccoides\u003c/em\u003e and \u003cem\u003eAegilops tauschii\u003c/em\u003e (Bd-Ta, Td-Ta and Aet-Ta) and wheat. These results suggest that \u003cem\u003eELP\u003c/em\u003e genes in wheat are distantly related to those in dicotyledonous species and are most closely associated with those in \u003cem\u003eAegilops tauschii\u003c/em\u003e, which might be because \u003cem\u003eAegilops tauschii\u003c/em\u003e are widely considered to be the D-genome ancestor of wheat [\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. The Ka/Ks ratio indicates the selection pressure of plant genes and can be used to diagnose the evolutionary form of the sequence [\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]. Thus, we calculated the Ka, Ks, Ka/Ks and T values of all orthologous gene pairs in wheat to further investigate the evolutionary trends of the \u003cem\u003eELP\u003c/em\u003e gene family (Table S2). The results showed that the Ka/Ks ratios of all orthologous genes (Ta-Gm, Ta-Os, Ta-Bd, Ta-Td, and Ta-Aet) were less than 1, suggesting that purification selection plays a dominant role in the evolutionary trend of the \u003cem\u003eELP\u003c/em\u003e gene family. According to the divergence time T value calculated from the Ks value, we found that the divergence time of the orthologous genes (Ta-Gm, Ta-Os, Ta-Bd, Ta-Td, and Ta-Aet) was different, among which the orthologous genes (Ta-Gm) had the longest divergence time and had the shortest divergence time with \u003cem\u003eAegilops tauschii\u003c/em\u003e. In conclusion, \u003cem\u003eTaELPs\u003c/em\u003e genes in wheat may have evolved from orthologous genes of other plant species or closely related plants.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e2.5. Gene structure and conserved motif analysis of \u003cem\u003eTaELPs\u003c/em\u003e genes\u003c/h2\u003e \u003cp\u003eThe phylogenetic tree constructed based on the protein sequences of the members of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family showed that the members of the \u003cem\u003eTaELPs\u003c/em\u003e gene family were divided into three groups (GroupA, GroupB and GroupC), and the results were visualized by combining the gene structure and conserved motifs (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The gene structure analysis of wheat \u003cem\u003eTaELPs\u003c/em\u003e found that introns ranged from 4\u0026ndash;10, and exons ranged from 5\u0026ndash;11. A maximum of 11 exons were found in the \u003cem\u003eTaELP1-A\u003c/em\u003e, while a minimum of five introns were found in the \u003cem\u003eTaELP5-A\u003c/em\u003e and \u003cem\u003eTaELP5-D\u003c/em\u003e. In the \u003cem\u003eTaELPs\u003c/em\u003e members of GroupA, GroupB and GroupC, the exon-intron numbers of genes were relatively close, and the exon-intron structure of most genes was relatively conservative. For example, in Group A, the number of introns and exons of \u003cem\u003eTaELPs\u003c/em\u003e genes were mostly 9 and 10, and the number of untranslated regions (UTRs) was relatively close.\u003c/p\u003e \u003cp\u003eTo further understand the structural diversity of wheat \u003cem\u003eELPs\u003c/em\u003e, we submitted the protein sequences of 18 \u003cem\u003eTaELPs\u003c/em\u003e genes to the MEME5.4.1 online website and predicted 10 conserved motifs (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eB). The results showed that the number of conserved motifs ranged from 3 to 9. All \u003cem\u003eTaELPs\u003c/em\u003e gene family members contained motif 2, while \u003cem\u003eTaELP4-A\u003c/em\u003e, \u003cem\u003eTaELP4-B\u003c/em\u003e, and \u003cem\u003eTaELP4-D\u003c/em\u003e lacked motif 1. Motif 5 was unique to \u003cem\u003eTaELP3-A\u003c/em\u003e, \u003cem\u003eTaELP3-B\u003c/em\u003e, and \u003cem\u003eTaELP3-D\u003c/em\u003e. The same group of \u003cem\u003eELP\u003c/em\u003e proteins contained similar motifs, and they may also have similarities in gene function. For example, Motif 2, Motif 8, Motif 4, Motif 10 and Motif 7 were included in Group A. The differences in the types and numbers of conserved motifs in wheat \u003cem\u003eELP\u003c/em\u003e proteins reflected the structural diversity of these proteins, indicating that they might have different biological functions.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e2.6. Protein conservation domain and 3-D protein structure analysis of \u003cem\u003eTaELPs\u003c/em\u003e gene\u003c/h2\u003e \u003cp\u003ePfam database was utilized to find the important component domains of \u003cem\u003eTaELPs\u003c/em\u003e proteins [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]. The conserved domains of \u003cem\u003eTaELPs\u003c/em\u003e were shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e. \u003cem\u003eTaELP1-A\u003c/em\u003e, \u003cem\u003eTaELP1-B\u003c/em\u003e, and \u003cem\u003eTaELP1-D\u003c/em\u003e contain four WD40(WD domain, G-beta repeat) protein domains; \u003cem\u003eTaELP4-A\u003c/em\u003e, \u003cem\u003eTaELP4-B\u003c/em\u003e and \u003cem\u003eTaELP4-D\u003c/em\u003e contain 1 Elong_Iki1 (Elongator subunit Iki1) domain; \u003cem\u003eTaELP2-A\u003c/em\u003e, \u003cem\u003eTaELP2-B\u003c/em\u003e and \u003cem\u003eTaELP2-D\u003c/em\u003e contain 1 IKI3 domain; \u003cem\u003eTaELP6-A\u003c/em\u003e, \u003cem\u003eTaELP6-B\u003c/em\u003e, and \u003cem\u003eTaELP6-D\u003c/em\u003e contain 1 PAXNEB domain; \u003cem\u003eTaELP3-A\u003c/em\u003e, \u003cem\u003eTaELP3-B\u003c/em\u003e and \u003cem\u003eTaELP3-D\u003c/em\u003e contain a catalytic domain of S-adenosylmethionine (Radical SAM superfamily) and a histone acetyltransferase (Acetyltransferase (GNAT) family) domain; \u003cem\u003eTaELP5-A\u003c/em\u003e, \u003cem\u003eTaELP5-B\u003c/em\u003e and \u003cem\u003eTaELP5-D\u003c/em\u003e all contain an \u003cem\u003eELP6\u003c/em\u003e (Elongator complex 6) domain, In addition, an Elong_Iki1 (Elongator subunit Iki1) domain was found in \u003cem\u003eTaELP5-B\u003c/em\u003e and \u003cem\u003eTaELP5-D\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eWe used SWISS-MODEL to further identify 3-D models of \u003cem\u003eTaELPs\u003c/em\u003e proteins[\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e] and the 3-D structure reveals a few key residues linked to biological processes or intended outcomes, (Figure S1). For \u003cem\u003eTaELP1-A\u003c/em\u003e, \u003cem\u003eTaELP1-B\u003c/em\u003e and \u003cem\u003eTaELP1-D\u003c/em\u003e proteins, 3-D structures were analyzed using the template \"6qk7.1.B\", a template that describes Elongator complex protein 2. \u003cem\u003eTaELP2-A\u003c/em\u003e, \u003cem\u003eTaELP2-B\u003c/em\u003e and \u003cem\u003eTaELP2-D\u003c/em\u003e proteins, 3-D structures were analyzed using the \"6qk7.1.A\" template, a template describing the Elongator complex protein 1. \u003cem\u003eTaELP3-A\u003c/em\u003e, \u003cem\u003eTaELP3-B\u003c/em\u003e, and \u003cem\u003eTaELP3-D\u003c/em\u003e Protein, the 3-D structure was analyzed using the \"6qk7.1.C\" template, a description of Elongator complex protein 3, and identified as containing two ligands (1 x 5AD and 1 x SF4), of which, 5AD (5'-DEOXYADENOSINE)9 residues within 4\u0026Aring; and 4 PLIP interactions, SF4(IRON/SULFUR CLUSTER)8 residues within 4\u0026Aring; and 3 PLIP interactions. \u003cem\u003eTaELP4-A\u003c/em\u003e, \u003cem\u003eTaELP4-B\u003c/em\u003e and \u003cem\u003eTaELP4-D\u003c/em\u003e, 3-D structures were analyzed using the\"4a8j.1.B\" template, \"4a8j.1.B\" template was a description ELONGATOR COMPLEX PROTEIN 5; \u003cem\u003eTaELP5-A\u003c/em\u003e used \"4ejs.1.C\" template to analyze 3-D structure, \u003cem\u003eTaELP5-B\u003c/em\u003e and \u003cem\u003eTaELP5-D\u003c/em\u003e proteins, used \"4wia.1.A\" to analyze 3-D structure; \u003cem\u003eTaELP6-A\u003c/em\u003e,\u003cem\u003eTaELP6-B\u003c/em\u003e and \u003cem\u003eTaELP6-D\u003c/em\u003e proteins, used The \"4a8j.1.A\" template analyzes 3-D structures. The Residues in the favored region of the Ramachandran plots generated by all \u003cem\u003eTaELPs\u003c/em\u003e ranged from 86.39% to 94.32; the Residues in the outlier region ranged from 0.44\u0026ndash;6.27%; Coverage of most \u003cem\u003eTaELPs\u003c/em\u003e was above 80%, only \u003cem\u003eTaELP4-A\u003c/em\u003e, \u003cem\u003eTaELP4- B\u003c/em\u003e and \u003cem\u003eTaELP4-D\u003c/em\u003e protein coverage was 58% (Table S5).\u003c/p\u003e \u003cp\u003eIn addition, we also used SOPMA to calculate the secondary structure elements of the protein sequence (Table S6), the results showed that the \u003cem\u003eTaELPs\u003c/em\u003e protein α-helix (Alpha helix) ranged from 13.52\u0026ndash;44.46%; β-turn (Beta turn) ranged from 3.39\u0026ndash;10.24%; Random coil ranged from 30.31\u0026ndash;49.94%; Extended strand ranged from 9.66\u0026ndash;32.30%.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e2.7. Cis-acting element regulation (CARE) analysis of wheat \u003cem\u003eTaELPs\u003c/em\u003e genes\u003c/h2\u003e \u003cp\u003eA total of 91 different CAREs were identified by analyzing the upstream 2000bp promoter region of the \u003cem\u003eTaELPs\u003c/em\u003e gene, mainly investigating abiotic stress and defense-related hormone response elements. All of the identified CAREs were divided into five groups according to their known functions (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eB, Table S7). Group I contained 48 environmental stress-related CAREs. Among them, 14 different types of abiotic stress response elements, one cis-element involved in low-temperature response (LTR), one cis-acting element (DRE core) regulating cold and dehydration response gene expression, three anaerobic-induced Essential cis-regulatory elements (ARE, GC-motif, plant_AP-2-like), one MYB binding site (MBS) associated with drought induction, and eight stress response-related response elements (such as MYC, as-1, Unnamed__1, WRE3, etc.) (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eA, Table S7/S8). 4 cis-acting elements related to wounding and pathogen response (box S, TC-rich repeats, W box, CCAAT-box), the rest were light-responsive elements of different types, such as 3-AF1 binding site, AE-box, Box II, GT1-motif, chs-CMA1a, etc.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eGroup II contained hormone response-related CAREs. There was a total of 13 different types of CAREs that regulate hormone response, such as cis-acting elements involved in abscisic acid response (ABRE, ABRE2), cis-regulatory elements involved in MeJA response (CGGTA-motif, TGACG-motif), cis-regulatory elements (AuxRR-core, TGA-element) involved in auxin response, and cis-acting elements (TCA-element, TGACG-motif) related to salicylic acid response, etc. Group III contained four core cis-acting elements, among which, CAAT-box and TATA-box appear most frequently in all \u003cem\u003eTaELPs\u003c/em\u003e genes, indicating that they play an important role in transcription initiation. TATA-box (including TATA and ATTATA-box) and CAAT-box cis-elements are promoter-associated elements that function at the initiation of transcription [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]. Group IV was plant growth and development-related CAREs, including cis-elements involved in seed-specific expression (AAGAA-motif, RY-element), cis-acting elements involved in cell cycle regulation (MSA-like), and meristem expression associated cis-regulatory elements (CAT-box) and several other CAREs associated with cell division. Group V was a small number of CAREs with unknown functions, which are also commonly found in the promoter sequences of \u003cem\u003eTaELPs\u003c/em\u003e genes, indicating that they may also be involved in the regulatory mechanism of \u003cem\u003eTaELPs\u003c/em\u003e genes on the environment (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eA, \u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eB, Table S7).\u003c/p\u003e \u003cp\u003eIn conclusion, \u003cem\u003eTaELPs\u003c/em\u003e genes may be involved in the regulation of the above environmental stress-related, phytohormone responses, and cell growth and development. These transcription factors CAREs play an important role to induce transcription of \u003cem\u003eTaELPs.\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e2.8. \u003cem\u003eTaELPs\u003c/em\u003e expression pattern prediction analysis\u003c/h2\u003e \u003cp\u003eTo further explore the expression patterns of \u003cem\u003eTaELPs\u003c/em\u003e genes in different tissues, developmental stages, and abiotic stresses in wheat, we retrieved all wheat mRNA transcription data from the wheat expression database and visualized TPM values with a heat map (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). The results showed that \u003cem\u003eTaELP3-A, TaELP3-B, TaELP3-D\u003c/em\u003e, and \u003cem\u003eTaELP5-D\u003c/em\u003e were expressed at high levels in the tissues of seedlings, roots, stems, leaves, and inflorescences at various stages. In addition, we found that \u003cem\u003eTaELP5-D\u003c/em\u003e was up-regulated in flag leaves with the prolongation of post-flowering time, while \u003cem\u003eTaELP3-A\u003c/em\u003e, \u003cem\u003eTaELP3-B\u003c/em\u003e, and \u003cem\u003eTaELP3-D\u003c/em\u003e were gradually down-regulated in flag leaves. Therefore, we speculated that TaELPs genes may be related to senescence; However, TaELP1-D and TaELP6-A had lower expression levels in all different tissues and developmental stages of wheat. \u003cem\u003eTaELP4-A\u003c/em\u003e、\u003cem\u003eTaELP4-B\u003c/em\u003e and \u003cem\u003eTaELP4-D\u003c/em\u003e were found to be at higher levels expressed in the root, stem, leaf, and spike tissues. \u003cem\u003eTaELP1-A\u003c/em\u003e, \u003cem\u003eTaELP1-B\u003c/em\u003e, \u003cem\u003eTaELP2-A\u003c/em\u003e, \u003cem\u003eTaELP2-B\u003c/em\u003e, \u003cem\u003eTaELP2-D\u003c/em\u003e, \u003cem\u003eTaELP5-A\u003c/em\u003e, \u003cem\u003eTaELP5-B\u003c/em\u003e, \u003cem\u003eTaELP6-B\u003c/em\u003e, and \u003cem\u003eTaELP6-D\u003c/em\u003e had tissue expression specificity in the stem, milk grain stage、stem 1 cm spike、leaf, seven leaf stage and roots, three-leaf stage.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cem\u003eTaELP3-A\u003c/em\u003e, \u003cem\u003eTaELP3-B\u003c/em\u003e, \u003cem\u003eTaELP3-D\u003c/em\u003e, and \u003cem\u003eTaELP5-D\u003c/em\u003e had the most obvious up-regulation of gene expression after drought treatment 6h (Figure S2); similarly, the gene expression was up-regulated most obviously after heat treatment 6h; After drought and heat stress treatment, gene expression was slightly down-regulated. \u003cem\u003eTaELP3-A\u003c/em\u003e, \u003cem\u003eTaELP3-B\u003c/em\u003e, and \u003cem\u003eTaELP3-D\u003c/em\u003e had higher expression in low-temperature stress. In salt stress, \u003cem\u003eTaELP3-A\u003c/em\u003e, \u003cem\u003eTaELP3-\u003c/em\u003eB, and \u003cem\u003eTaELP3-D\u003c/em\u003e were all up-regulated to varying degrees and showed a downward trend as a whole; \u003cem\u003eTaELP4-A\u003c/em\u003e, \u003cem\u003eTaELP4-B\u003c/em\u003e, and \u003cem\u003eTaELP4-D\u003c/em\u003e had the highest up-regulated expression after salt stress 48h and showed a trend of upward. \u003cem\u003eTaELP5-D\u003c/em\u003e showed very low expression in different treatment times of salt stress; the expression patterns of \u003cem\u003eTaELP2-B\u003c/em\u003e and \u003cem\u003eTaELP1-D\u003c/em\u003e genes showed an upward trend and the up-regulated expression was most obvious at salt stress 48h; The gene expression of other \u003cem\u003eTaELPs\u003c/em\u003e genes were slightly up-regulated or down-regulated in different treatment times of salt stress (Figure S2).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e2.9. Expression pattern validation analysis of \u003cem\u003eTaELPs\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eFurther understand the potential response mechanism of wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family in tolerance to abiotic stress, hormone response, and leaf senescence, we detected the transcription patterns of all \u003cem\u003eTaELPs\u003c/em\u003e genes in abiotic stresses (drought, salt, and dark treatment), hormones treatments (IAA, SA, ABA), and during leaf senescence. Under drought treatment, \u003cem\u003eTaELP2\u003c/em\u003e exhibited down-regulated transcriptions at most of the time points. The expression of other \u003cem\u003eTaELPs\u003c/em\u003e genes was upregulated to varying degrees at different times of drought treatment. Among them, \u003cem\u003eTaELP3\u003c/em\u003e, \u003cem\u003eTaELP1\u003c/em\u003e, and \u003cem\u003eTaELP4\u003c/em\u003e were significantly upregulated, And \u003cem\u003eTaELP3\u003c/em\u003e significant upregulation was observed after 6h and 72h of drought treatment, The overall trend of TaELP1was upregulated, and significant upregulation was observed from 24h to 48h. The expression of \u003cem\u003eTaELP4\u003c/em\u003e was most significantly upregulated after 12h of drought treatment (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eUnder salt stress, \u003cem\u003eTaELP3\u003c/em\u003e, \u003cem\u003eTaELP1\u003c/em\u003e, and \u003cem\u003eTaELP6\u003c/em\u003e were slightly up-regulated or down-regulated compared with the control (0h); TaELP4 was significantly down-regulated under salt stress 6h, 12h, 24h, and 48h, but after 72h of treatment, The expression was significantly up-regulated immediately; The expression of \u003cem\u003eTaELP2\u003c/em\u003e and \u003cem\u003eTaELP5\u003c/em\u003e was up-regulated, \u003cem\u003eTaELP2\u003c/em\u003e were significantly upregulated from 48h to 72h, And \u003cem\u003eTaELP5\u003c/em\u003e significant upregulation was observed after 6h and 72h of salt treatment (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e).To explore the induced expression patterns of all members of the \u003cem\u003eTaELPs\u003c/em\u003e gene family in plant growth and development, we performed dark treatments at different times (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e). The results showed that the expression of all \u003cem\u003eTaELPs\u003c/em\u003e genes was up-regulated to varying degrees in the early or late stage of dark treatment. For example, the expression pattern of the TaELP2 gene showed an up-regulated trend; \u003cem\u003eTaELP3\u003c/em\u003e, \u003cem\u003eTaELP1\u003c/em\u003e, \u003cem\u003eTaELP4\u003c/em\u003e, \u003cem\u003eTaELP6\u003c/em\u003e genes showed an up-regulated trend from rising to decline; \u003cem\u003eTaELP5\u003c/em\u003e significant upregulation was observed after 72h of dark treatment (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eUnder IAA treatment, the expression patterns of most \u003cem\u003eTaELPs\u003c/em\u003e genes were down-regulated, and only \u003cem\u003eTaELP3\u003c/em\u003e and \u003cem\u003eTaELP5\u003c/em\u003e genes were up-regulated. \u003cem\u003eTaELP3\u003c/em\u003e was most significantly up-regulated after 48h of IAA treatment; \u003cem\u003eTaELP5\u003c/em\u003e was up-regulated immediately after 6h of \u003cem\u003eIAA\u003c/em\u003e treatment and the expression level reached a peak and then showed a downward trend (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eA). Under SA treatment, it was observed that only \u003cem\u003eTaELP5\u003c/em\u003e exhibited low expression levels at all time treatment stages and other \u003cem\u003eTaELPs\u003c/em\u003e genes were up-regulated to varying degrees. For example, \u003cem\u003eTaELP2\u003c/em\u003e was significantly up-regulated in the early and late stages of SA treatment; the gene expression patterns of \u003cem\u003eTaELP4\u003c/em\u003e and \u003cem\u003eTaELP6\u003c/em\u003e were significantly up-regulated after 12h of SA treatment; \u003cem\u003eTaELP3\u003c/em\u003e and \u003cem\u003eTaELP1\u003c/em\u003e significant upregulation was observed after 24h of SA treatment (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eB).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eUnder ABA treatment, the relative expression levels of all \u003cem\u003eTaELPs\u003c/em\u003e genes were significantly different (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eC). \u003cem\u003eTaELP6\u003c/em\u003e showed a significant upregulation at all-time treatment stages compared to the control. \u003cem\u003eTaELP1\u003c/em\u003e, \u003cem\u003eTaELP2\u003c/em\u003e, \u003cem\u003eTaELP4\u003c/em\u003e, and \u003cem\u003eTaELP5\u003c/em\u003e showed an overall upward trend, and the gene expression was up-regulated most significantly after 72h of treatment. Moreover, \u003cem\u003eTaELP3\u003c/em\u003e showed a significant upregulation only at ABA treatment 6h compared to the control (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eC).\u003c/p\u003e \u003cp\u003eDuring leaf senescence, all \u003cem\u003eTaELPs\u003c/em\u003e genes were up-regulated to varying degrees in the late senescence (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e). The up-regulation trend of \u003cem\u003eTaELP2\u003c/em\u003e and \u003cem\u003eTaELP6\u003c/em\u003e is the same, and the overall showed a trend from decline to rise, \u003cem\u003eTaELP2\u003c/em\u003e was most significantly expressed at 30 days after flowering, followed by \u003cem\u003eTaELP6\u003c/em\u003e at 10 days after flowering; \u003cem\u003eTaELP5\u003c/em\u003e was significantly up-regulated at 10 days and 19 days after flowering; \u003cem\u003eTaELP1\u003c/em\u003e, \u003cem\u003eTaELP3\u003c/em\u003e, and \u003cem\u003eTaELP4\u003c/em\u003e were up-regulated in the same trend. Overall, members of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family play important regulatory roles in abiotic stresses, hormones, and leaf senescence.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e2.10. Prediction of protein-protein interactions of wheat ELPs\u003c/h2\u003e \u003cp\u003eTo study the interaction between wheat \u003cem\u003eTaELPs\u003c/em\u003e and other proteins, a network was constructed using the STRING database (Figure S3, Table S9). Based on the predicted results, we observed that TaELP1, TaELP2, TaELP3 and TaELP6 had protein interactions with a Chromatin associated protein KTI12 (Traes_5BL_92F800E16.1, Traes_5BL_D8ECD483D.2 and Traes_5DL_A9A62BF38.1) and a Diphthamide biosynthesis protein 3 (Traes_7BL_69CD9E49D.2, Traes_7DL_EF5C1F9EA.1). In addition, TaELP3 had protein interactions with Traes_2AS_03ED0D137.1, Traes_2BS_E0BE8F2D1.1 and Traes_2DS_E75C5D4AC.1 which encodes WD40 repeat-containing proteins. No protein interacting with \u003cem\u003eTaELP4\u003c/em\u003e and \u003cem\u003eTaELP5\u003c/em\u003e was found.\u003c/p\u003e \u003cp\u003eWD repeats proteins are widely present in eukaryotes and are involved in various cellular behavioral and physiological regulations, such as signal transduction, activation of transcriptional activity, cell growth and development, and control of apoptosis. The presence of WD40 domains or repeated WD40 motifs can act as a scaffold for protein-protein or protein-DNA assembly, play an important role in protein interactions and can act as a mediator of transient interactions between other proteins [\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]. Besides, chromatin-associated protein KTI12 was found to interact with the Elongator complex (ELP) in the process of RNA polymerase II promoting transcription elongation [\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eClimate warming is a common challenge for global agricultural development and improving agricultural biomass production. According to the predictions of climate change models, global crop yields have been declining as the climate changes, but the decrease in wheat yields is closely related to abiotic stresses [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e]. One of the effective ways to deal with the current bottleneck of wheat production is to tap more wheat stress resistance gene resources and apply them to the breeding of wheat resistant varieties. Elongator complex (ELP) is an indispensable component of gene transcription in eukaryotes, and can also indirectly participate in cell behaviors such as extracellular secretion, telomere gene silencing and DNA damage repair by modifying tRNA to participate in the translation process, which influenced abiotic and biotic stress responses as well as plant growth and development [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e].\u003c/p\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e4.1. Evolution and genetic relationship of \u003cem\u003eTaELPs\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eIn higher plants, there was a gap between the number of \u003cem\u003eELP\u003c/em\u003e gene family members in closely genetic relationship plants. A total of 6 \u003cem\u003eELP\u003c/em\u003e genes were identified in \u003cem\u003eArabidopsis\u003c/em\u003e, and whereas, 6 members were identified in rice [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e], which is more closely related to wheat and fewer numbers that were identified in wheat. In the process of studying plant evolution, it is found that the expansion of gene families is closely related to the occurrence of gene duplication, which may originate from segmental, tandem or whole-genome duplication [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e]. Segment duplication (SD) is common in biological evolution. When the repetitive DNA sequence exceeds 1kbp or the identity is higher than 90%, we consider that the gene has SD on the chromosome [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e]. In this study, it was found that the members of the wheat TaELPs gene family contained 18 pairs of \u003cem\u003eELP\u003c/em\u003e paralogous genes, all of which originated from large segment duplication. There were two fragment duplications were observed on 2A, 2B, 2D, 4A, 4B, and 4D and one fragment duplication was observed on 1A, 1B, 1D, 7A, 7B, and 7D (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eD, Table S2). These results suggest that fragment duplication may have a dominant role in the evolution and expansion of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family. The evolutionary selection pressure (Ka/Ks) and divergence time (MYA) of 18 pairs of \u003cem\u003eTaELPs\u003c/em\u003e genes were calculated, and the Ka/Ks\u0026thinsp;\u0026lt;\u0026thinsp;1 of all duplicated gene pairs indicated that \u003cem\u003eTaELPs\u003c/em\u003e belonged to purifying selection in long-term evolution. The phylogeny and cluster analysis of wheat \u003cem\u003eTaELPs\u003c/em\u003e protein showed that wheat ELP protein was more closely related to the evolution of monocotyledonous plants.\u003c/p\u003e \u003cp\u003eIn addition, to further understand the genetic relationship of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family, we investigated the collinearity relationship of \u003cem\u003eELPs\u003c/em\u003e genes in wheat and other species (including monocotyledonous and dicotyledonous plants). No wheat orthologous gene pairs were found in \u003cem\u003eArabidopsis\u003c/em\u003e, only 3 \u003cem\u003eELPs\u003c/em\u003e gene orthologous gene pairs were found in Glycine max, and most wheat orthologous gene pairs were obtained in \u003cem\u003eOryza sativa\u003c/em\u003e, \u003cem\u003eBrachypodium distachyon\u003c/em\u003e, \u003cem\u003eTriticum dicoccoides\u003c/em\u003e, and \u003cem\u003eAegilops tauschii\u003c/em\u003e. It has been shown that allohexaploid bread wheat (AABBDD) is produced by crossing the allotetraploid \u003cem\u003eTriticum dicoccoides\u003c/em\u003e (BBAA) with the diploid \u003cem\u003eAegilops tauschii\u003c/em\u003e (DD) containing the D genome [\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e]. Therefore, we speculated that hybridization was part of the reason for more orthologous gene pairs between wheat and \u003cem\u003eTriticum dicoccoides\u003c/em\u003e and \u003cem\u003eAegilops tauschii\u003c/em\u003e. The 56 orthologous gene pairs identified in wheat were all derived from whole-genome duplication or fragment duplication within the genome. We also calculated evolutionary selection pressure (Ka/Ks) and divergence time (MYA) between wheat and other species, with Ka/Ks\u0026thinsp;\u0026lt;\u0026thinsp;1 for all duplicate gene pairs, suggesting that \u003cem\u003eTaELPs\u003c/em\u003e belonged to purification selection in long-term evolution. The results of this study suggest that \u003cem\u003eTaELPs\u003c/em\u003e genes in wheat may have evolved from orthologous genes in other plant species and are more closely related to monocotyledonous plants.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003e4.2. Structural diversity of wheat \u003cem\u003eTaELPs\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eThe protein conserved domain analysis showed (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e) that each \u003cem\u003eELP\u003c/em\u003e subunit protein in wheat contains its corresponding conserved domain, which further illustrated the evolutionary and functional conservation of eukaryotic \u003cem\u003eELPs\u003c/em\u003e [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e]. Compared with the previously reported domains of \u003cem\u003eArabidopsis\u003c/em\u003e and rice \u003cem\u003eELPs\u003c/em\u003e and wheat TaELPs, we found that there were significant differences in the number of domains between some subunits, such as \u003cem\u003eOsELP2\u003c/em\u003e and \u003cem\u003eAtELP2\u003c/em\u003e contained 5 and 6 WD40 repeat protein domains, respectively, \u003cem\u003eOsELP6\u003c/em\u003e and \u003cem\u003eAtELP6\u003c/em\u003e contained 2 and 1 Elongation complex protein 6 domains respectively[\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e], while wheat contained 1 Elongation complex protein 6 domain and 4 WD40 repeat protein domains, the size of the domains were similar, and the overall difference was not large. It showed that in different plant species, there were slight differences between the structure and number of \u003cem\u003eELPs\u003c/em\u003e, suggesting that there may be changes in function. The WD40 repeat protein domain contained in \u003cem\u003eELP2\u003c/em\u003e has been widely reported to participate in a variety of biological processes of plant growth and development, and play an important role in protein-protein and protein-DNA interactions. Plant anthocyanin biosynthesis and abiotic stress response are closely related.[\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]. IKI3 was a chromatin-associated domain that interacts with ELP and contains the WD40 repeat protein during RNA polymerase II-promoted transcription elongation [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]. ELP6 was an accessory subunit of the Elongator complex, was associated with histone acetylation in the nucleus and tRNA modification in the cytoplasm and was able to catalyze the elongation of transcription by RNAII [\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e]. PAXNEB has been found in different eukaryotes. It is a component of RNA polymerase II extension protein subunit and HAP subcomplex, which can catalyze intracellular histone acetylation [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e53\u003c/span\u003e]. Members of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family had different domains, suggesting that the six subunits had different potential functions in plants.\u003c/p\u003e \u003cp\u003eThe gene structure and conserved motif analysis of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene showed (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e) that there were quantitative differences in the exon-intron number of the \u003cem\u003eTaELPs\u003c/em\u003e gene, which was similar to the results in \u003cem\u003eArabidopsis\u003c/em\u003e and rice[\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]. Among the 10 identified motifs, all members of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family contained a conserved motif of MOTIF2. These results suggested that the evolutionary pattern of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family was relatively conserved. Further, subcellular localization prediction showed that most members of the wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family were located in the cytoplasm, and a few were located in the nucleus, plastid and plasma membrane. Subcellular localization results of the Elongator complex have been reported in a variety of plants. Nelissen and colleagues (2010) detected GFP-ELO3 fusion protein in the nucleus by the GFP fusion protein method [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e]. Two years later, Tran et al. found ELP3 red fluorescent protein under the detection of confocal laser scanning microscopy through the epidermal cells of faba bean leaves, and the results again showed that the Elongator complex exists in the nucleus and cytoplasm [\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e55\u003c/span\u003e]. In conclusion, most of the above reports have confirmed that the elongator complex (ELP) is localized in the nucleus and cytoplasm, which is also confirmed by our prediction of the subcellular localization of all \u003cem\u003eTaELPs\u003c/em\u003e genes in wheat.\u003c/p\u003e \u003cp\u003e \u003cspan\u003e \u003cp\u003e \u003cb\u003e4.3 Transcription analysis of\u003c/b\u003e \u003cspan type=\"BoldItalic\" class=\"BoldItalic\" name=\"Emphasis\"\u003eTaELP\u003c/span\u003e \u003cb\u003egenes reveals its role in wheat growth, development and abiotic stress tolerance\u003c/b\u003e\u003c/p\u003e\u003c/span\u003e \u003c/p\u003e \u003cp\u003eDifferent CAREs distribution in promoter regions may indicate variations in gene regulation and function [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e56\u003c/span\u003e]. Through the analysis of 2000 bp cis-acting elements upstream of the promoter of wheat \u003cem\u003eTaELPs\u003c/em\u003e gene family members, we can further understand the process of \u003cem\u003eTaELPs\u003c/em\u003e gene-regulating physiological changes in wheat. We found that all TaELPs genes contain more than one cis-acting element in response to abiotic stress. The frequency of drought-induced cis-elements MYC, MYB, MBS, etc, and stress defense response cis-elements STRE, as-1, as-1, ARE, etc, and other stress-related CAREs in all TaELPs genes (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eB, Table S8) very high [\u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e56\u003c/span\u003e]. MYB and MYC transcription factor binding elements were involved in plant responses to drought, high salt, and low temperature, and regulate the expression of related genes under stress[\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e57\u003c/span\u003e, \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e58\u003c/span\u003e]. In the study of drought resistance of various crops, it was found that MBS elements generally exist in the promoter sequences of drought resistance-related genes and are closely related to the response to drought [\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e59\u003c/span\u003e] indicating that they play an important role in abiotic stress. In addition to the above abiotic stress response elements, they also contained defense and stress response elements and a variety of biological stress-related transcription factor binding elements. We also found hormone-related elements, including ABRE (abscisic acid-responsive), TCA-element (salicylic acid-responsive), CGTCA-motif (MeJA responsive), p-box, and TATC-box (gibberellin responsive element), etc. ABRE response element was related to ABA-related gene expression, which was regulated by ABA-dependent or ABA-independent in abiotic stress [\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e60\u003c/span\u003e]. indicating that \u003cem\u003eTaELPs\u003c/em\u003e may be involved in signaling during the wheat stress response. In addition, the frequency of many cells cycle-related CAREs and CAREs with unknown functions was also high, indicating that \u003cem\u003eELPs\u003c/em\u003e genes in wheat have different functions during wheat development. The cis-acting elements of \u003cem\u003eELP\u003c/em\u003e family genes were closely related to their possible physiological processes, Predicting the cis-acting elements of unknown gene families is helpful to quickly predict gene functions, It is an effective way to screen candidate genes and analyze gene functions by reverse genetics [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e, \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e61\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe tissue expression patterns of genes are often closely related to their gene functions. The 18 \u003cem\u003eTaELPs\u003c/em\u003e genes were expressed to varying degrees in each stage of seedling, root, stem, leaf and panicle, indicating that they may play an important role in the growth and development of wheat plants [\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e]. When plants encounter drought stress, the root system will initiate a stress response mechanism to cope with drought by reducing water evaporation and root growth [\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e63\u003c/span\u003e, \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e64\u003c/span\u003e]. The published transcriptome data show that \u003cem\u003eTaELP1\u003c/em\u003e, \u003cem\u003eTaELP3\u003c/em\u003e, and \u003cem\u003eTaELP4\u003c/em\u003e were expressed at high levels in roots, and their transcription levels were significantly increased under drought stress, suggesting that \u003cem\u003eTaELP1\u003c/em\u003e, \u003cem\u003eTaELP3\u003c/em\u003e, and \u003cem\u003eTaELP4\u003c/em\u003e may affect the root development of plants and respond to drought stress are closely related. qRT-PCR results showed that the expression levels of \u003cem\u003eTaELP1\u003c/em\u003e, \u003cem\u003eTaELP3\u003c/em\u003e, and \u003cem\u003eTaELP4\u003c/em\u003e were significantly increased under drought stress, indicating that the above \u003cem\u003eTaELPs\u003c/em\u003e genes play an important role in the regulatory mechanism of plants resisting drought stress (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e). Soil salinization is a limiting factor for today's agricultural development. When studying salt-tolerant plants, it was found that the root system is very sensitive to salt stress and can quickly initiate a stress response mechanism [\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e, \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e]. The results of qRT-PCR showed that the expression levels of \u003cem\u003eTaELP2\u003c/em\u003e and \u003cem\u003eTaELP4\u003c/em\u003e were significantly increased during 48\u0026ndash;72 h of salt stress, and the expression levels of \u003cem\u003eTaELP5\u003c/em\u003e were significantly up-regulated in the early and late stages of salt stress (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e). Transcriptome analysis showed that \u003cem\u003eTaELP2\u003c/em\u003e, \u003cem\u003eTaELP4\u003c/em\u003e and \u003cem\u003eTaELP5\u003c/em\u003e were highly expressed in root tissue, and their transcription levels were significantly increased in the late stage of salt stress, indicating that \u003cem\u003eTaELP2\u003c/em\u003e, \u003cem\u003eTaELP4\u003c/em\u003e and \u003cem\u003eTaELP5\u003c/em\u003e genes play an important role in the regulation mechanism of plant salt stress resistance. Studies have shown that Elongator plays an important role in plant growth and development. In the phenotypic observation of Arabidopsis elo/Atelp mutants, it was found that the germination and seed setting rates at the seedling stage were lower, the germination was slower, the flowering was delayed, the morphological structure of leaves and flowers was abnormally developed, and the leaf area was reduced, The apical meristem develops irregularly, and the underground taproot and hypocotyl grow slowly [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e].This study found that \u003cem\u003eTaELP3\u003c/em\u003e, \u003cem\u003eTaELP4\u003c/em\u003e, \u003cem\u003eTaELP5\u003c/em\u003e and \u003cem\u003eTaELP6\u003c/em\u003e were highly expressed in flag leaves at different time points after flowering (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e), and their transcription levels were significantly increased under dark stress(Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e), qRT-PCR results showed that all \u003cem\u003eTaELPs\u003c/em\u003e genes were up-regulated to varying degrees in flag leaves (24 to 30 days after flowering), and \u003cem\u003eTaELP6\u003c/em\u003e was the most significantly up-regulated, suggesting that \u003cem\u003eTaELP3, TaELP4\u003c/em\u003e, \u003cem\u003eTaELP5\u003c/em\u003e are closely related to wheat senescence and \u003cem\u003eTaELP6\u003c/em\u003e may play an important role in wheat senescence (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCell proliferation is closely related to Elongator, and cell proliferation is often induced by plant hormones [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e67\u003c/span\u003e]. We performed 3 hormone treatments (IAA, SA and ABA) on all \u003cem\u003eTaELPs\u003c/em\u003e in wheat. When studying the expression of ELP-related genes in a variety of plants, it was found that a large number of auxin genes have high differential expression, so we speculate that Elongator may induce the expression of auxin-related genes to control plant growth and development [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e]. Further studies found that auxin-related genes were hypoacetylated at histone H3K14, suggesting that the Elongator complex may interact with RNAPII to catalyze the formation and transcription of chromatin, thereby promoting the expression of auxin-related genes [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e, \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e]. Therefore, we treated all \u003cem\u003eTaELPs\u003c/em\u003e with IAA at different times and found that only \u003cem\u003eTaELP3\u003c/em\u003e had a higher expression, suggesting that \u003cem\u003eTaELP3\u003c/em\u003e may be an important regulator of wheat growth and development (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eA). Salicylic acid (SA) was a signaling molecule that initiates stress response mechanisms when plants encounter pathogens [\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e68\u003c/span\u003e], SA accumulates after pathogen infection and was critical for activating local and systemic acquired resistance [\u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e69\u003c/span\u003e]. Elongator inhibits the expression of CAT3 and other related antioxidant genes and can promote the expression of SA-related genes [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]. In all \u003cem\u003eTaELPs\u003c/em\u003e treated with SA at different times, all \u003cem\u003eTaELPs\u003c/em\u003e except \u003cem\u003eTaELP5\u003c/em\u003e were expressed and \u003cem\u003eTaELP2\u003c/em\u003e was significantly expressed, indicating that wheat \u003cem\u003eTaELP2\u003c/em\u003e may play an irreplaceable role in pathogen defense (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eB). Under ABA stress treatment, the expression of \u003cem\u003eTaELP3\u003c/em\u003e was not significantly up-regulated, and the remaining \u003cem\u003eTaELPs\u003c/em\u003e genes were up-regulated to varying degrees under ABA treatment at different times (Fig.\u0026nbsp;\u003cspan refid=\"Fig10\" class=\"InternalRef\"\u003e10\u003c/span\u003eC). We found that \u003cem\u003eTaELP2\u003c/em\u003e, \u003cem\u003eTaELP4\u003c/em\u003e, \u003cem\u003eTaELP5\u003c/em\u003e and TaELP6 were significantly up-regulated in the early and late stages of ABA stress treatment. Plant hormone ABA can cause plant cell behavior regulation and physiological signal transduction, and affect different stages of plant growth and development. When subjected to abiotic stresses such as drought, salt, and high temperature, ABA can conduct signals to activate plant stress response mechanisms [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e, \u003cspan additionalcitationids=\"CR71 CR72\" citationid=\"CR70\" class=\"CitationRef\"\u003e70\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e73\u003c/span\u003e]. According to related reports, ABA signaling molecules were involved in RNA-related processes such as RNA splicing, RNA structure stabilization, and RNA elongation, indicating that ABA signaling was closely related to RNA metabolism and regulation [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e, \u003cspan additionalcitationids=\"CR75 CR76 CR77\" citationid=\"CR74\" class=\"CitationRef\"\u003e74\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e78\u003c/span\u003e]. ABA signaling was associated with the histone acetyltransferase Elongator complex [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e]. Therefore, we analyzed the relationship between ABA stress and the three abiotic stresses. The results showed that the expression levels of \u003cem\u003eTaELP2\u003c/em\u003e and \u003cem\u003eTaELP4\u003c/em\u003e were significantly increased at 48-72h induced by salt stress and ABA, and the expression levels of \u003cem\u003eTaELP5\u003c/em\u003e were significantly increased at 6h and 72h. indicating that \u003cem\u003eTaELP2\u003c/em\u003e, \u003cem\u003eTaELP4\u003c/em\u003e and \u003cem\u003eTaELP5\u003c/em\u003e genes play an important role in the regulatory mechanism of plants resisting salt stress and may exist in the salt stress response signaling pathway dependent on ABA regulation. All the above results showed that \u003cem\u003eTaELP2\u003c/em\u003e, \u003cem\u003eTaELP3\u003c/em\u003e, \u003cem\u003eTaELP4\u003c/em\u003e, and \u003cem\u003eTaELP6\u003c/em\u003e may be important regulators of abiotic stress and leaf senescence in wheat, and also play an important role in signal transduction. Therefore, this temporal and spatial expression pattern of \u003cem\u003eTaELP\u003c/em\u003e genes indicates that these \u003cem\u003eELPs\u003c/em\u003e might have a function in different tissues and various developmental stages as well as abiotic stress tolerance in wheat.\u003c/p\u003e \u003cp\u003eThrough predictive analysis of interactions between wheat TaELPs and other proteins (Figure S3, Table S9), We found that TaELPs interacting proteins play roles in plant growth and development, hormonal and abiotic stress responses. The three interacting proteins contained the WD40 domain. The WD-repeat protein can be used as a scaffold for protein-protein assembly, and may interact with TaELPs to play a role in plants, which can regulate plant growth and development, transcriptional regulation, hormone signaling, and initiate plant stress defense mechanisms [\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e, \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e]. The WD40 repeat protein had been extensively studied in \u003cem\u003eArabidopsis\u003c/em\u003e, and it can act as a regulator of plant growth and development to regulate plant-specific development [\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e]. Chromatin-associated protein KTI12 is an important regulator of the Elongator complex and is involved in the modification of uridine bases in eukaryotic tRNA, and there is a close physical and functional relationship between them [\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e, \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e80\u003c/span\u003e]. Two stress-related genes PtKTI12A and PtKTI12B were identified in \u003cem\u003ePopulus trichocarpa\u003c/em\u003e under high temperature and drought stress and their expression levels were analyzed in each specific tissue, and it was found that they were differentially expressed [\u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e81\u003c/span\u003e]. The results showed that in the stress response mechanism, KTI12 protein can be induced to express, activate plant resistance to stress and participate in tRNA swing uridine modification [\u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e81\u003c/span\u003e]. Thus, TaELPs along with their interacting partners might be required to develop wheat stress tolerance. Overall, these findings are useful in elucidating the specific biological activities of \u003cem\u003eTaELP\u003c/em\u003e genes in order to generate high-yielding and stress-tolerant wheat cultivars.\u003c/p\u003e \u003c/div\u003e"},{"header":"5. Conclusions","content":"\u003cp\u003eWheat is a major grain crop and a staple meal all over the world. Therefore, researchers have intended to improve wheat production, quality, and different stress tolerance. The current study was utilized the biological information methods to identify \u003cem\u003eELP\u003c/em\u003e family genes in wheat and carried out a comprehensive and systematic analysis. The gene structure, amino acid motif, and subcellular localization prediction results of the \u003cem\u003eTaELPs\u003c/em\u003e gene family showed that the \u003cem\u003eELP\u003c/em\u003e gene in wheat is highly conserved. The homology analysis between wheat and other plant species showed that the \u003cem\u003eTaELPs\u003c/em\u003e gene had a close homology relationship with \u003cem\u003eAegilops tauschii\u003c/em\u003e. \u003cem\u003eCis\u003c/em\u003e-acting element analyzed predicts 14 different types of abiotic stress response elements and 13 hormone response-related CAREs. Transcriptome data combined with qRT-PCR results showed that The \u003cem\u003eTaELP\u003c/em\u003e genes have a predominant role in the regulatory mechanism of wheat development and stress tolerance. The findings of this work will aid in understanding the role of ELPs in plant developmental processes and various stress situations, as well as their sequential implementation to boost yield and generate stress-tolerant wheat cultivars.\u003c/p\u003e"},{"header":"6. Materials And Methods","content":"\u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003e6.1. Identification of wheat ELPs family gene members\u003c/h2\u003e \u003cp\u003eWe used 12 \u003cem\u003eELP\u003c/em\u003e genes from \u003cem\u003eArabidopsis\u003c/em\u003e and rice to find members of the \u003cem\u003eELP\u003c/em\u003e gene family in the wheat genome [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e]. The ELP protein sequences of \u003cem\u003eArabidopsis\u003c/em\u003e and rice were retrieved from the Ensemble Plants database. ELP genes were identified from the whole wheat genome by BLASTp against the most recent wheat entire genes from IWGSC (RefSeq v1.0) (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://plants.ensembl.org/index.html\u003c/span\u003e\u003cspan address=\"http://plants.ensembl.org/index.html\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e, E-value\u0026thinsp;\u0026lt;\u0026thinsp;1e\u003csup\u003e-5\u003c/sup\u003e and bit-score\u0026thinsp;\u0026gt;\u0026thinsp;100) and through the Blast Compare Two Seqs tool of TBtools [\u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e82\u003c/span\u003e]. Finally, after eliminating duplicated sequences, the output of BLASTp and TBtools were selected for domain analysis. SMART (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://smart.embl-heidelberg.de/\u003c/span\u003e\u003cspan address=\"http://smart.embl-heidelberg.de/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) or InterPro (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.ebi.ac.uk/interpro\u003c/span\u003e\u003cspan address=\"https://www.ebi.ac.uk/interpro\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) or NCBI CDD (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://ncbi.nlm.gov.Structure/cdd/cdd.shtml\u003c/span\u003e\u003cspan address=\"https://ncbi.nlm.gov.Structure/cdd/cdd.shtml\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and HMM scan (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.ebi.ac.uk/Tools/hmmer/search/hmmscan\u003c/span\u003e\u003cspan address=\"https://www.ebi.ac.uk/Tools/hmmer/search/hmmscan\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) were utilized to check for the existence of \u003cem\u003eELP\u003c/em\u003e gene family domains in the remaining sequences. The length, molecular weight, isoelectric point (pI), and gross average value (GRAVY) of the wheat ELP proteins were calculated using ProtParam software (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://web.expasy.org/protparam/\u003c/span\u003e\u003cspan address=\"https://web.expasy.org/protparam/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003e6.2. Sequence alignment and phylogenetic tree construction\u003c/h2\u003e \u003cp\u003eFull-length protein sequences from several species were aligned using ClustalW in MEGA X [\u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e83\u003c/span\u003e], and all sequences after alignment analysis were imported into MEGA X for constructing a phylogenetic tree using the neighbor-joining (NJ) method [\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e84\u003c/span\u003e] with 1000 bootstrap values [\u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e85\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003e6.3. Chromosomal localization, gene duplication and collinearity analysis\u003c/h2\u003e \u003cp\u003eFor chromosomal distribution, \u003cem\u003eELPs\u003c/em\u003e genomic positions were obtained from the EnsemblPlants BioMart (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://plants.ensembl.org/biomart/martview/\u003c/span\u003e\u003cspan address=\"http://plants.ensembl.org/biomart/martview/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). The \u003cem\u003eELPs\u003c/em\u003e were numbered according to their ascending chromosomal location and were given a \u0026ldquo;Ta\u0026rdquo; prefix. TBtools was used to visualize the \u003cem\u003eTaELPs\u003c/em\u003e on the wheat chromosomes. McscanX software was used to investigate tandem and segmental duplications within the \u003cem\u003eTaELP\u003c/em\u003e gene family and collinearity between \u003cem\u003eELPs\u003c/em\u003e from wheat and several other species [\u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e86\u003c/span\u003e]. The TBtools were used to compute the non-synonymous substitution rate (Ka), synonymous substitution rate (Ks), and the Ka/Ks ratio and the divergence time T was estimated by T\u0026thinsp;=\u0026thinsp;Ks/(2\u0026times;9.1\u0026times;10^-5)Mya [\u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003e6.4. Subcellular localization and 3-D Structure Modeling\u003c/h2\u003e \u003cp\u003eSubcellular localization was predicted using WoLF PSORT (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://wolfpsort.hgc.jp/\u003c/span\u003e\u003cspan address=\"https://wolfpsort.hgc.jp/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and CELLO v.2.5(\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://cello.life.nctu.edu.tw\u003c/span\u003e\u003cspan address=\"http://cello.life.nctu.edu.tw\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). The SWISS-MODEL (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://swissmodel.expasy.org/interactive#sequence\u003c/span\u003e\u003cspan address=\"https://swissmodel.expasy.org/interactive#sequence\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) was used to create three-dimensional (3D) protein structure of TaELPs.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003e6.5. Structure, domain and motif analysis of TaELP genes\u003c/h2\u003e \u003cp\u003eWheat genome annotation file (GFF3 format), CDS sequence and protein sequence of \u003cem\u003eELP\u003c/em\u003e gene were retrieved from the Ensemble Plants database (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://plants.ensembl.org/index.html\u003c/span\u003e\u003cspan address=\"http://plants.ensembl.org/index.html\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). The gene-related information of \u003cem\u003eTaELPs\u003c/em\u003e was extracted using the function of Gtf/Gff3 Sequences Extract of TBtools [\u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e]. The Gene Structure Display Server 2.0 (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://gsds.gao-lab.org/\u003c/span\u003e\u003cspan address=\"http://gsds.gao-lab.org/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) was used to visualize intron, exon, and untranslated regions. The ELP gene domains were retrieved from the Pfam database and Evolview (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.evolgenius.info/evolview-v2/\u003c/span\u003e\u003cspan address=\"https://www.evolgenius.info/evolview-v2/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) was used to visualize the gene domains. To anticipate the conserved motifs of TaELPs, we employed the motif-based sequence analysis tools MEME version 5.4.1 (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://meme-suite.org/tools/meme\u003c/span\u003e\u003cspan address=\"http://meme-suite.org/tools/meme\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) with a maximum section of up to 10 motifs and visualized using TBtools [\u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003e6.6. Analysis of cis-acting regulatory elements (CAREs) and protein interaction networks\u003c/h2\u003e \u003cp\u003eThe 2000 bp upstream sequences of 18 \u003cem\u003eTaELPs\u003c/em\u003e genes were retrieved from the Ensemble Plants database (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://plants.ensembl.org/index.html\u003c/span\u003e\u003cspan address=\"http://plants.ensembl.org/index.html\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) and submitted to the online software PlantCARE (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://bioinformatics.psb.ugent.be/webtools/plantcare/html/\u003c/span\u003e\u003cspan address=\"http://bioinformatics.psb.ugent.be/webtools/plantcare/html/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) to find CAREs. Subsequently, the TBtools Heatmap was used for data visualization [\u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e]. The STRING online server (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://string-db.org/cgi\u003c/span\u003e\u003cspan address=\"https://string-db.org/cgi\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) was used to predict the protein interaction network of TaELPs.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec23\" class=\"Section2\"\u003e \u003ch2\u003e6.7. Tissue specific expression and stress analysis of wheat TaELPs\u003c/h2\u003e \u003cp\u003eThe wheat (Chinese spring) RNA-Seq data was retrieved from the Wheat Exp database (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://wheat-expression.com/\u003c/span\u003e\u003cspan address=\"http://wheat-expression.com/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), and the TPM (transcripts per million reads) value was used to evaluate the transcript abundance of wheat \u003cem\u003eTaELPs\u003c/em\u003e genes. The MeV tool was used to visualize the expression [\u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e88\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec24\" class=\"Section2\"\u003e \u003ch2\u003e6.8. Plant Materials and treatments\u003c/h2\u003e \u003cp\u003eTwo winter wheat varieties were used in this study, including Yannong 19 (registration no. 2004003) and Jinmai 39 (registration no. ZM0218364). These varieties have been approved by the Ministry of Agriculture and Rural Affairs of the People's Republic of China and are widely cultivated in the traditional farming systems of Shandong and Shanxi provinces. Yannong 19 and Jinmai 39 seeds were maintained and provided by the Yantai Academy of Agricultural Sciences (Shandong, China) and the Institute of Economic Crops of Shanxi Agricultural University (Shanxi Academy of Agricultural Sciences) (Shanxi, China), respectively.\u003c/p\u003e \u003cp\u003eWheat cultivar, Jinmai 39 was selected to study the expression of \u003cem\u003eTaELPs\u003c/em\u003e under abiotic stress (drought stress, salt stress, dark stress) and hormone treatments (ABA, IAA, SA). The Seedlings were grown in an artificial climate box (16h light/8h dark) at 22\u0026deg;C. When the wheat plants grew in the 2\u0026ndash;3 leaf stage, they were treated with sterile ddH\u003csub\u003e2\u003c/sub\u003eO (control) or 20% PEG-6000 and 250 mM NaCl solutions, respectively. For abscisic acid (ABA), auxin (IAA), salicylic acid (SA) treatments, wheat plants at the same stage were sprayed with 100 mM ABA, IAA, SA, and 0.1% (v/v) ethanol (control). Yannong 19, a delayed senescence wheat cultivar was cultivated in the field and flag leaf samples were taken at 0, 7, 16, 19, 22, 24, 25, and 30 days after anthesis. All leaves collected were immediately frozen in liquid nitrogen and stored in a -80\u0026deg;C freezer for further RNA extraction. Each of the above experiments was set up with 3 sets of repetitions.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec25\" class=\"Section2\"\u003e \u003ch2\u003e6.9. RNA extraction, cDNA first-strand synthesis and real-time PCR analysis\u003c/h2\u003e \u003cp\u003eThe Quick RNA isolation Kit (Tiangen Biochemical Technology Co., Ltd.) was used to extract RNA according to the manufacturer\u0026rsquo;s instructions and DNase I treatment was used to remove DNA contamination. Synthesis of the first strand of cDNA was carried out according to the instructions of the kit (Baori Doctor Biotechnology (Beijing) Co., Ltd.). To measure the expression of \u003cem\u003eTaELPs\u003c/em\u003e, qRT-PCR analysis was performed with specific primers (Table S1). The \u003cem\u003eElongation factor 1a\u003c/em\u003e (\u003cem\u003eTaEF-1a\u003c/em\u003e) was used as an internal reference gene (GenBank accession no. Q03033) [\u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e89\u003c/span\u003e]. The threshold values (CT) were generated using the ABI PRISM 7500 system (Applied Biosystems, Foster City, CA, USA), and the transcription level of \u003cem\u003eTaELPs\u003c/em\u003e were assessed using the comparative 2\u003csup\u003e-ΔΔCT\u003c/sup\u003e method [\u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e90\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e"},{"header":"Abbreviations","content":"\u003cp\u003eELP:\u0026nbsp;Elongator complexes; RNAPII: RNA polymerase II; CAREs: \u003cem\u003eCis\u003c/em\u003e-acting regulatory elements; qRT-PCR: Quantitative real-time PCR; CTD: C-terminal repeat domain; Fe-S: iron-sulfur radical; SAM: S-adenosylmethionine; HAT: histone acetyltransferase; SAR: systemic acquired resistance; ORF: open reading frame; PI: predicted isoelectric point; Ka: nonsynonymous mutation rate; Ks: synonymous mutation rate; ABA: abscisic acid; IAA: auxin; SA: salicylic acid.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research was sponsored by the State Key Laboratory of Sustainable Dryland Agriculture, Shanxi Agricultural University (No. 202002-2); Shanxi Agricultural University Academic Recovery Special Project (2020xshf02) and Supported by Scientific and Technological Innovation Programs of Higher Education Institutions in Shanxi (2021L124).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor information\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAffiliations\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFeng Guo, Md Ashraful Islam, Xiujuan Jin, Lili Sun, Kai Zhao, Juan Lu, Rongyue Yan, Ning Li, Shuguang Wang and Daizhen Sun.\u003c/p\u003e\n\u003cp\u003eState Key Laboratory of Sustainable Dryland Agriculture, College of Agronomy, Shanxi Agricultural University, Taigu 030801, China.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFG designed the experiment. DZS and MAI provide advice on experimental protocols. XJJ, LLS, KZ, JL and RYY conducted experiments. FG wrote the manuscript. The manuscript was revised by DZS and MAI. The manuscript was grammatically revised and polished by MAI. All authors have read and agree to the published version of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCorresponding authors\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eCorrespondence to Daizhen Sun.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe have obtained the permissions to collect wheat cultivars, Yannong 19 and Jinmai 39 which were acquired from Yantai Academy of Agricultural Sciences (Shandong, China) and the Institute of Economic Crops of Shanxi Agricultural University (Shanxi, China), respectively and identified by Dr. Daizhen Sun. Jinmai 39 was grown in the greenhouse of Shanxi Agricultural University (SXAU). Yannong 19 was grown in the State Key Laboratory of Sustainable Dryland Agriculture, Shanxi Agricultural University. The research conducted in this study required neither the approval of the ethics committee nor any human or animal subjects. The described field studies do not require specific permission. The site is not privately owned or protected in any way, and field research does not involve endangered or protected species. The collection and experimental research of wheat materials involved in this study were approved by Shanxi Agricultural University (SXAU) in China and comply with the relevant guidelines and regulations.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll needed genome sequences and genome annotation files of wheat were obtained from \u003cem\u003eEnsemblplants\u003c/em\u003e database (https://plants.ensembl.org/Triticum_aestivum/Info/Index). The transcriptome sequencing data of different tissues and various abiotic stresses used in this study were retrieved from the Wheat Exp database (http://wheat-expression.com/) and these RNA-sequencing reads of the Wheat Exp database were previously deposited with NCBI under accession codes PRJEB25639, PRJEB23056, PRJNA436817, SRP133837, PRJEB25640, and PRJEB25593. All data generated in this study are available in public and also included in the article and its Additional files. \u003c/p\u003e\n\u003cp\u003eIn addition, all databases used in this study are open in public and the links are as follows:\u003c/p\u003e\n\u003cp\u003eEnsemblPlants: https://plants.ensembl.org/Triticum_aestivum/Info/Index \u003c/p\u003e\n\u003cp\u003eExPasy: http://web.expasy.org/compute_ pi/ \u003c/p\u003e\n\u003cp\u003eSTRING: https://cn.string-db.org/cgi/input?sessionId=bXZTbla6vfa4 \u003c/p\u003e\n\u003cp\u003e Wheat Expression Browser: http://www.wheat-expression.com/ \u003c/p\u003e\n\u003cp\u003eWheat eFP Browser: http://bar.utoronto.ca/efp_wheat/cgi-bin/efpWeb.cgi \u003c/p\u003e\n\u003cp\u003eGSDS: http://gsds.cbi.pku.edu.cn/ \u003c/p\u003e\n\u003cp\u003eMEME: http://meme-suite.org/ \u003c/p\u003e\n\u003cp\u003eplantCARE: http://bioinformatics.psb.ugent.be/webtools/plantcare/html/ \u003c/p\u003e\n\u003cp\u003eSMART: http://smart.embl-heidelberg.de\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no conflict of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eBradsher J, Jackson K, Conaway R, Conaway J: \u003cstrong\u003eRNA polymerase II transcription factor SIII. 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In: \u003cem\u003eBiomedical Informatics for Cancer Research.\u003c/em\u003e 2010: 267-277.\u003c/li\u003e\n\u003cli\u003ePaolacci A, Tanzarella O, Porceddu E, Ciaffi M: \u003cstrong\u003eIdentification and validation of reference genes for quantitative RT-PCR normalization in wheat\u003c/strong\u003e. \u003cem\u003eBMC molecular biology \u003c/em\u003e2009, \u003cstrong\u003e10\u003c/strong\u003e:11.\u003c/li\u003e\n\u003cli\u003eChang S, Chen W, Yang J: \u003cstrong\u003eAnother formula for calculating the gene change rate in real-time RT-PCR\u003c/strong\u003e. \u003cem\u003eMolecular biology reports \u003c/em\u003e2009, \u003cstrong\u003e36\u003c/strong\u003e(8):2165-2168.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Elongator complexes (ELPs), In silico analysis, abiotic stress, leaf senescence, wheat","lastPublishedDoi":"10.21203/rs.3.rs-1521902/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-1521902/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cstrong\u003eBackground:\u003c/strong\u003e Elongator complexes (ELPs) are the protein complexes that promote transcription through histone acetylation in eukaryotic cells and interact with elongating RNA polymerase II (RNAPII). ELPs role in plant growth and development, signal transduction, and response to biotic and abiotic stresses have been confirmed in model plants. However, the functions of the wheat \u003cem\u003eELP\u003c/em\u003e genes are not well documented. \u003c/p\u003e\u003cp\u003e\u003cstrong\u003eResults:\u003c/strong\u003e The present study was identified 18 members of the \u003cem\u003eELPs\u003c/em\u003e from the wheat genome by a homology search and further bioinformatics and transcriptions patterns in response to different stress conditions were analyzed to dissect their potential regulatory mechanisms in wheat. Gene duplication analysis showed that 18 pairs of \u003cem\u003eELP\u003c/em\u003e paralogous genes were derived from segmental duplication, which was divided into 6 clades by protein phylogenetic and cluster analysis. The orthologous analysis of wheat \u003cem\u003eTaELPs\u003c/em\u003e genes showed that \u003cem\u003eTaELP\u003c/em\u003e genes may have evolved from orthologous genes of other plant species or closely related plants. Moreover, a variety of \u003cem\u003eCis\u003c/em\u003e-acting regulatory elements (CAREs) related to growth and development, hormone response, biotic and abiotic stresses were identified in the \u003cem\u003eTaELPs \u003c/em\u003epromoter region. Publicly available RNA-seq data analysis indicated that \u003cem\u003eTaELPs\u003c/em\u003e gene family members were differentially expressed in wheat seedlings, roots, stems, and leaf panicles, as well as under abiotic stresses. Further, the qRT-PCR analysis showed that the transcription of \u003cem\u003eTaELPs\u003c/em\u003e was induced under hormone, salt, and drought stress and during leaf senescence. \u003c/p\u003e\u003cp\u003e\u003cstrong\u003eConclusions:\u003c/strong\u003e Overall, \u003cem\u003eTaELP \u003c/em\u003egenes might be regulated by hormone signaling pathways and responded to abiotic stress and leaf senescence, which could be investigated further as a potential candidate gene for wheat abiotic stress tolerance and yield improvement.\u003c/p\u003e","manuscriptTitle":"Insights into the bioinformatics and transcriptional analysis of the Elongator complexes (ELPs) Gene Family of wheat: TaELPs contribute to wheat abiotic stress tolerance and leaf senescence","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2022-07-22 17:04:01","doi":"10.21203/rs.3.rs-1521902/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"0a0a5b78-b6ff-4b81-88c4-8da2e6b218de","owner":[],"postedDate":"July 22nd, 2022","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2022-09-13T07:14:21+00:00","versionOfRecord":[],"versionCreatedAt":"2022-07-22 17:04:01","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-1521902","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-1521902","identity":"rs-1521902","version":["v1"]},"buildId":"WvIrzKhiLBfengagbw6Ux","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}