Transcription Factors In Wheat Research Articles

Genome-wide identification and functional characterization of wheat Brassinazole-resistant transcription factors in response to abiotic stresses and stripe rust infection.

Brassinazole-resistant (BZR) transcription factors (TFs) are key players in brassinolides (BRs) signaling pathway, which is widely involved in regulating plant growth and development, as well as in plant responding to a variety stresses. Despite their critical roles, little is known about BZR TFs in wheat. In this study, we performed genome-wide analysis of BZR gene family from wheat genome, and 20 TaBZRs were identified. Based on the phylogenetic relationships of TaBZR and BZRs from rice and Arabidopsis, all BZR genes were clustered into four groups. The intron-exon structural patterns and conserved protein motifs of TaBZRs showed high group specificity. TaBZR5, 7, and 9 were significantly induced after salt, drought treatment, and stripe rust infection. However, TaBZR16, which was significantly upregulated under NaCl application, was not expressed during wheat-stripe rust fungus interaction. These results indicated that BZR genes in wheat play different roles in response to various stresses. The results of this study will lay a foundation for further in-depth functional studies of TaBZRs and will provide information for the breeding and genetic improvement of wheat against drought and salt stresses.

TaABI19 positively regulates grain development in wheat

Starch is the most important component in the endosperm, and its synthesis is regulated by multiple transcription factors (TFs) in cereals. However, whether the functions of these TFs are conserved among cereals remains unclear. In this study, we cloned a B3 family TF in wheat, named TaABI19, based on its orthologous sequence in maize (Zea mays L.). Alignment of the DNA and protein sequences showed that ABI19 is conserved in maize and wheat (Triticum aestivum L.). We found that TaABI19 is highly expressed in young spikes and developing grains, and encodes a nucleus-localized transcriptional activator in wheat. The taabi19-b1 null mutants obtained by EMS exhibited a down-regulation of starch synthesis, shorter grain length and lower thousand-grain weight (TGW). Furthermore, we proved that TaABI19 could bind to the promoters of TaPBF homologous genes and enhance their expression. Haplotype association showed that TaABI19-B1 is significantly associated with TGW. We found that Hap2 and Hap3 were favored and had undergone positive selection in China’s wheat breeding programs. Less than 50% of the modern cultivars convey the favored haplotypes, indicating that TaABI19 still can be considered as a target locus for marker-assisted selection breeding to increase TGW in China.

Transcription factor retention through multiple polyploidization steps in wheat.

Whole-genome duplication is widespread in plant evolutionary history and is followed by nonrandom gene loss to return to a diploid state. Across multiple angiosperm species, the retained genes tend to be dosage-sensitive regulatory genes such as transcription factors, yet data for younger polyploid species is sparse. Here, we analyzed the retention, expression, and genetic variation in transcription factors in the recent allohexaploid bread wheat (Triticum aestivum L.). By comparing diploid, tetraploid, and hexaploid wheat, we found that, following each of two hybridization and whole-genome duplication events, the proportion of transcription factors in the genome increased. Transcription factors were preferentially retained over other genes as homoeologous groups in tetraploid and hexaploid wheat. Across cultivars, transcription factor homoeologs contained fewer deleterious missense mutations than nontranscription factors, suggesting that transcription factors are maintained as three functional homoeologs in hexaploid wheat populations. Transcription factor homoeologs were more strongly coexpressed than nontranscription factors, indicating conservation of function between homoeologs. We found that the B3, MADS-M-type, and NAC transcription factor families were less likely to have three homoeologs present than other families, which was associated with low expression levels and high levels of tandem duplication. Together, our results show that transcription factors are preferentially retained in polyploid wheat genomes although there is variation between families. Knocking out one transcription factor homoeolog to alter gene dosage, using TILLING or CRISPR, could generate new phenotypes for wheat breeding.

Identification and expression profiles of the YABBY transcription factors in wheat.

BackgroundYABBY is a plant-specific transcription factor (TF) that belongs to the zinc finger protein superfamily and is composed of a C2–C2 domain at the N-terminus and a YABBY domain at the C-terminus. It plays a role in plant development and growth.MethodsIn this study, 20 YABBY TFs were identified in the wheat genome. Phylogenetic relationships, collinearity relationships, gene structures, conserved motifs, and expression patterns were analyzed.ResultsTwenty TaYABBY TFs were distributed unevenly on 15 chromosomes. Collinearity analysis showed that these genes have a close relationship with monocot plants. The phylogenetic tree of wheat YABBYs classified these TaYABBYs into FIL, YAB2, INO, and CRC clades. Gene structure and conserved motif analyses showed that they share similar components in the same clades. Expression profile analysis showed that many TaYABBY genes have high expression levels in leaf tissues and are regulated by abiotic stresses, especially salt stress. Our results provide a basis for further functional characterization of the YABBY gene family.

RAV transcription factor regulatory function in response to salt stress in two Iranian wheat landraces

Amongst the transcription factor groups, the AP2/ERF (Apetala2/Ethylene Response Factor) superfamily is one of the main groups in plants and plays an essential role in tolerating abiotic and biotic stresses. The AP2/ERF superfamily consists of ERF, AP2, RAV, and Soloist families based on the AP2 domain number. The RAV (Related to ABI3/VP1) family members have been revealed to be stimulate by a number of biotic and abiotic environmental incentives; including pathogen infection, salicylic acid, osmotic stress, cold, high salinity, wounding, and exogenous hormone application. However, limited data are available on the contributions of RAV transcription factors in wheat (Triticum aestivum L.). In the present study, a total of 26 RAV genes were identified in wheat from a genome-wide search against the latest wheat genome data. Phylogenetic and sequence alignment analyses divided the wheat RAV genes into 4 clusters, I, II, III and IV. Chromosomal distribution, gene structure and motif composition were subsequently investigated. The 26 TaRAV genes were unevenly distributed on 21 chromosomes. After cloning and sequencing of 7 TaRAVs candidate genes the expression levels of two TaRAVs, TaRAV4 and TaRAV5, were validated through qPCR analyses in two salt-tolerant Iranian landraces of wheat. Our results showed that the TaRAV4 and TaRAV5 were co-expressed in wheat tissues and were highly correlated to salt tolerance indices such as the K+/Na+ ratio. Protein interaction revealed that the TaRAV4 and TaRAV5 were related to vital proteins such as PK4 and PP2C, and MYB and Zinc finger transcription factors, and Gigantea proteins. This study improved our knowledge of the RAV gene family function in wheat and the probable role of RAVs in salt tolerance mechanisms to improve crop production under changing environments. Also, the two relatively salt-tolerant landraces of wheat that were examined in this study could be suitable candidates for future breeding studies.

IN SILICO GENE EXPRESSION ANALYSIS OF THE STRESS-RELATED NAC-A GENE SUBFAMILY TO DISSECT THEIR ROLE IN ABIOTIC STRESS TOLERANCE IN BREAD WHEAT (TRITICUM AESTIVUM L.)

Wheat is a major staple crop that is largely affected by different abiotic stresses that include heat, drought, and salinity. The main objective of this study was to identify wheat NAC transcription factors that are related to the NAC-a subfamily, which is involved in mediating stress tolerance in different plant species. Furthermore, in silico gene expression analysis was conducted to detect differential changes in wheat NAC-a subfamily members in different organs, developmental stages, and under various abiotic stress. Herein, using phylogenetic analysis for 488 NAC transcription factors, 41 proteins were identified as wheat NAC-a subfamily members. In silico gene expression analysis found that NAC-related wheat transcription factors are expressed exclusively at the anthesis stage till dough development with high expression levels detected in flag leaves. The in-silico gene expression analysis identified SNAC1-related members, which had high expression levels under drought, cold, and heat stresses. The identified stress-induced wheat NAC-a subfamily members can be utilized in the future to develop climate-smart wheat cultivars with improved tolerance against abiotic stresses.

Study of Expression Pattern of Some Transcription Factors in Wheat under Drought Stress and Zinc Nanoparticles

Study of Expression Pattern of Some Transcription Factors in Wheat under Drought Stress and Zinc Nanoparticles

Evolutionary Divergence and Biased Expression of NAC Transcription Factors in Hexaploid Bread Wheat (Triticum aestivum L.).

The NAC genes, a large plant-specific family of transcription factors, regulate a wide range of pathways involved in development and response to biotic and abiotic stress. In this study, the NAC transcription factors were identified in 27 green plants, and the results showed that NAC transcription factors in plants undergo an appearance stage from water to land and a number expansion stage from gymnosperm to angiosperm. Investigating the evolutionary process of the NAC transcription factors from diploid species to hexaploid wheat revealed that tandem replications during the polyploidization process is an important event for increasing the number of NAC transcription factors in wheat. Then, the molecular characteristics, phylogenetic relationships, and expression patterns of 462 NAC transcription factors of hexaploid wheat (TaNACs) were analyzed. The protein structure results showed that TaNAC was relatively conservative at the N-terminal that contains five subdomains. All these TaNACs were divided into Group I and Group II by phylogenetic analysis, and the TaNACs in Group I should undergo strong artificial selection based on single nucleotide polymorphism (SNP) analysis. Through genome synteny and phylogenetic analysis, these TaNACs were classified into 88 groups and 9 clusters. The biased expression results of these TaNACs showed that there are 24 groups and 67 groups of neofunctionalization genes under biotic and abiotic stress, respectively, and 16 groups and 59 groups of subfunctionalization genes. This shows that neofunctionalization plays an important role in coping with different stresses. Our study provides new insights into the evolution of NAC transcription factors in hexaploid wheat.

TaNAC48 positively regulates drought tolerance and ABA responses in wheat (Triticum aestivum L.)

NAC family transcription factors (TFs) are important regulators in plant development and stress responses. However, the biological functions of NAC TFs in wheat are rarely studied. In this study, 43 putative drought-induced NAC genes were identified from de novo transcriptome sequencing data of wheat following drought treatment. Twelve wheat NACs along with ten known stress-related NACs from Arabidopsis and rice were clustered into Group II based on a phylogenetic analysis. TaNAC48, which showed a higher and constitutive expression level in Group II, was selected for further investigation. TaNAC48 transcript was up-regulated by drought, PEG, H2O2 and abscisic acid (ABA) treatment and encoded a nuclear localized protein. Overexpression of TaNAC48 significantly promoted drought tolerance with increased proline content, and decreased rates of water loss, malondialdehyde (MDA), H2O2 and O2− content. Root length and a stomatal aperture assay confirmed that TaNAC48-overexpression plants increased sensitivity to ABA. Electrophoretic mobility shift assay (EMSA) and luciferase reporter analysis indicated that TaAREB3 could bind to a cis-acting ABA-responsive element (ABRE) on TaNAC48 promoter and activate the expression of TaNAC48. These results suggest that TaNAC48 is essential in mediating crosstalk between the ABA signaling pathway and drought stress responses in wheat.

Identification and Characterization of PLATZ Transcription Factors in Wheat.

The PLATZ (plant AT-rich protein and zinc-binding protein) transcription factor family is a class of plant-specific zinc-dependent DNA-binding proteins. PLATZ has essential roles in seed endosperm development, as well as promoting cell proliferation duration in the earlier stages of the crops. In the present study, 62 TaPLATZ genes were identified from the wheat genome, and they were unequally distributed on 15 chromosomes. According to the phylogenetic analysis, 62 TaPLATZ genes were classified into six groups, including two groups that were unique in wheat. Members in the same groups shared similar exon-intron structures. The polyploidization, together with genome duplication of wheat, plays a crucial role in the expansion of the TaPLATZs family. Transcriptome data indicated a distinct divergence expression pattern of TaPLATZ genes that could be clustered into four modules. The TaPLATZs in Module b possessed a seed-specific expression pattern and displayed obvious high expression in the earlier development stage of seeds. Subcellular localization data of TaPLATZs suggesting that they likely perform a function as a conventional transcription factor. This study provides insight into understanding the structure divergence, evolutionary features, expression profiles, and potential function of PLATZ in wheat.

Genome-wide identification and expression profiling of the SPL family genes in wheat

The SQUAMOSA promoter-binding protein-like (SPL) proteins constitute a family of plant-specific transcription factors that play a vital role in plant development. Wheat (Triticum aestivum, AABBDD) is universally well-known as a cash crop; however, the SPLs of this important crop have not been systematically investigated as yet. In this study, we conducted a genome-wide survey in wheat and found 56 SPL genes belonging to 19 homologous groups. The SPL genes were divided into seven classes using phylogenetic tree analyses. We mapped these genes onto the wheat chromosomes and examined their structures and conserved motifs. Moreover, we performed a synteny analysis on wheat, and summarized the SPL family as well as the evolutionary relationships between SPLs. Thereafter, we compared the expression patterns of wheat SPLs under different conditions, confirming that SPLs play an important role in spike development. To conclude, the SPLs in triplets have analogous structures and similar expression patterns. The three-pair triplet response to jasmonic acid (JA) and abscisic acid (ABA) was determined by quantitative real-time polymerase chain reaction (qRT-PCR). This work provides a comprehensive understanding of the SPL gene family in wheat. Our investigation of the wheat SPL gene family provides a starting point for additional functional studies of these significant transcription factors in wheat.

The Wheat GENIE3 Network Provides Biologically-Relevant Information in Polyploid Wheat.

Gene regulatory networks are powerful tools which facilitate hypothesis generation and candidate gene discovery. However, the extent to which the network predictions are biologically relevant is often unclear. Recently a GENIE3 network which predicted targets of wheat transcription factors was produced. Here we used an independent RNA-Seq dataset to test the predictions of the wheat GENIE3 network for the senescence-regulating transcription factor NAM-A1 (TraesCS6A02G108300). We re-analyzed the RNA-Seq data against the RefSeqv1.0 genome and identified a set of differentially expressed genes (DEGs) between the wild-type and nam-a1 mutant which recapitulated the known role of NAM-A1 in senescence and nutrient remobilisation. We found that the GENIE3-predicted target genes of NAM-A1 overlap significantly with the DEGs, more than would be expected by chance. Based on high levels of overlap between GENIE3-predicted target genes and the DEGs, we identified candidate senescence regulators. We then explored genome-wide trends in the network related to polyploidy and found that only homeologous transcription factors are likely to share predicted targets in common. However, homeologs which vary in expression levels across tissues are less likely to share predicted targets than those that do not, suggesting that they may be more likely to act in distinct pathways. This work demonstrates that the wheat GENIE3 network can provide biologically-relevant predictions of transcription factor targets, which can be used for candidate gene prediction and for global analyses of transcription factor function. The GENIE3 network has now been integrated into the KnetMiner web application, facilitating its use in future studies.

CRISPR/Cas9 editing of wheat TaQ genes alters spike morphogenesis and grain threshability

The TaQ alleles as one of the AP2-like transcription factors in common wheat (Triticum aestivum) play an important role in the evolution of spike characteristics from wild and domesticated emmer to modern wheat cultivars. Its loss-of-function mutant not only changed threshability and spike architecture but also affected plant height, flowering time, and floret structure. However, the comprehensive functions of TaAQ and TaDq genes in wheat have not been fully elucidated yet. Here, CRISPR/SpCas9 was used to edit wheat TaAQ and TaDq. We obtained homozygous plants in the T1 generation with loss of function of only TaAQ or TaDq and simultaneous loss of function of TaAQ and TaDq to analyze the effect of these genes on wheat spikes and floret shapes. The results demonstrated that the TaAQ-edited plants and the TaAQ and TaDq simultaneously-edited plants were nearly similar in spike architecture, whereas the TaDq-edited plants were different from the wild-type ones only in plant height. Moreover, the TaAQ-edited plants or the TaAQ and TaDq simultaneously-edited plants were more brittle than the wild-type and the TaDq-edited plants. Based on the expression profiling, we postulated that the VRN1, FUL2, SEP2, SEP5, and SEP6 genes might affect the number of spikelets and florets per spike in wheat by regulating the expression of TaQ. Combining the results of this report and previous reports, we conceived a regulatory network of wheat traits, including plant height, spike shape, and floral organs, which were influenced by AP2-like family genes. The results achieved in this study will help us to understand the regulating mechanisms of TaAQ and TaDq alleles on wheat floral organs and inflorescence development.

Overexpression of Maize ZmC1 and ZmR Transcription Factors in Wheat Regulates Anthocyanin Biosynthesis in a Tissue-Specific Manner.

Maize ZmC1 and ZmR transcription factors belong to the MYB-type and bHLH families, respectively, and control anthocyanin biosynthesis. In this study, Agrobacterium-mediated transformation was used to generate transgenic wheat plants that overexpress ZmC1 and ZmR or both, with the objective of developing anthocyanin-enriched wheat germplasm. Three kinds of stable transgenic wheat lines were obtained. The integration of target genes in the transgenic wheat plants was confirmed by fluorescence in situ hybridization (FISH) analysis. We found that single overexpression of ZmC1 regulates pigmentation in the vegetative tissues such as coleoptiles, auricles, and stems. The single overexpression of ZmR controls the coloration in reproductive tissue like spikelets and seeds. The simultaneous overexpression of ZmC1 and ZmR showed the strongest pigmentation in almost all tissues. Furthermore, quantitative real-time PCR (qRT-PCR) analysis revealed that expression of the two transgenes, and of two conserved homologous and six associated structural genes involved in anthocyanin biosynthesis in wheat were greatly up-regulated in the transgenic plants. Similarly, quantitative analysis for anthocyanin amounts based on HPLC-MS also confirmed that the transgenic wheat plants with combined overexpression of ZmC1 and ZmR accumulated the highest quantity of pigment products. Moreover, developing seeds overexpressing ZmR exposed to light conditions showed up-regulated transcript levels of anthocyanin biosynthesis-related genes compared to dark exposure, which suggests an important role of light in regulating anthocyanin biosynthesis. This study provides a foundation for breeding wheat materials with high anthocyanin accumulation and understanding the mechanism of anthocyanin biosynthesis in wheat.

Genome-Wide Identification and Analysis of the AP2 Transcription Factor Gene Family in Wheat (Triticum aestivum L.).

The AP2 transcription factors play important roles in regulating plant growth and development. However, limited data are available on the contributions of AP2 transcription factors in wheat (Triticum aestivum L.). In the present study, a total of 62 AP2 genes were identified in wheat from a genome-wide search against the latest wheat genome data. Phylogenetic and sequence alignment analyses divided the wheat AP2 genes into 3 clusters, euAP2, euANT, and basalANT. Chromosomal distribution, gene structure and duplication, and motif composition were subsequently investigated. The 62 TaAP2 genes were unevenly distributed on 21 chromosomes. Twenty-four homologous gene sets among A, B, and D sub-genomes were detected, which contributed to the expansion of the wheat AP2 gene family. The expression levels of TaAP2 genes were examined using the WheatExp database; most detected genes exhibited tissue-specific expression patterns. The transcript levels of 9 randomly selected TaAP2 genes were validated through qPCR analyses. Overexpression of TaAP2-10-5D, the most likely homolog of Arabidopsis ANT gene, increased organ sizes in Arabidopsis. Our results extend our knowledge of the AP2 gene family in wheat, and contribute to further functional characterization of AP2s during wheat development with the ultimate goal of improving crop production.

Genome-Wide Characterization and Expression Profiling of Squamosa Promoter Binding Protein-Like (SBP) Transcription Factors in Wheat (Triticum aestivum L.)

Transcription factors (TFs) play fundamental roles in the developmental processes of all living organisms. Squamosa Promoter Binding Protein-like (SBP/SBP-Box) is a major family of plant-specific TFs, which plays important roles in multiple processes involving plant growth and development. While some work has been done, there is a lot more that is yet to be discovered in the hexaploid wheat SBP (TaSBP) family. With the completion of whole genome sequencing, genome-wide analysis of SBPs in common hexaploid wheat is now possible. In this study, we used protein–protein Basic Local Alignment Search Tool (BLASTp) to hunt the newly released reference genome sequence of hexaploid wheat (Chinese spring). Seventy-four TaSBP proteins (belonging to 56 genes) were identified and clustered into five groups. Gene structure and motif analysis indicated that most TaSBPs have relatively conserved exon–intron arrangements and motif composition. Analysis of transcriptional data showed that many TaSBP genes responded to some biological and abiotic stresses with different expression patterns. Moreover, three TaSBP genes were generally expressed in the majority of tissues throughout the wheat growth and also responded to many environmental biotic and abiotic stresses. Collectively, the detailed analyses presented here will help in understanding the roles of the TaSBP and also provide a reference for the further study of its biological function in wheat.

Characteristics and Expression Pattern of MYC Genes in Triticum aestivum, Oryza sativa, and Brachypodium distachyon.

Myelocytomatosis oncogenes (MYC) transcription factors (TFs) belong to basic helix-loop-helix (bHLH) TF family and have a special bHLH_MYC_N domain in the N-terminal region. Presently, there is no detailed and systematic analysis of MYC TFs in wheat, rice, and Brachypodium distachyon. In this study, 26 TaMYC, 7 OsMYC, and 7 BdMYC TFs were identified and their features were characterized. Firstly, they contain a JAZ interaction domain (JID) and a putative transcriptional activation domain (TAD) in the bHLH_MYC_N region and a BhlH region in the C-terminal region. In some cases, the bHLH region is followed by a leucine zipper region; secondly, they display tissue-specific expression patterns: wheat MYC genes are mainly expressed in leaves, rice MYC genes are highly expressed in stems, and B. distachyon MYC genes are mainly expressed in inflorescences. In addition, three types of cis-elements, including plant development/growth-related, hormone-related, and abiotic stresses-related were identified in different MYC gene promoters. In combination with the previous studies, these results indicate that MYC TFs mainly function in growth and development, as well as in response to stresses. This study laid a foundation for the further functional elucidation of MYC genes.

Genome-wide Analysis of bZIP Transcription Factors in wheat and Functional Characterization of a TabZIP under Abiotic Stress

The basic leucine zipper (bZIP) represents one of the largest as well as most diverse transcription factor (TFs) families. They are known to play role in both stress as well as in various plant developmental processes. In the present study, a total of 191 bZIP transcription factors have been identified from Triticum aestivum. Expression analysis during various stress conditions, developmental stages, different varieties and gene ontology enrichment analysis suggest their possible roles in abiotic stress as well as in developmental responses. In the current analysis, one of the members named as TabZIP (Traes_7AL_25850F96F.1) was selected for detailed analysis to understand its role under different abiotic stress conditions. Gene expression studies revealed differential expression of TabZIP in various abiotic stress conditions like heat, salinity and dehydration suggesting the possible role of bZIP in various stress mitigation mechanism. Arabidopsis transgenics overexpressing TabZIP showed enhanced tolerance to salinity, drought, heat and oxidative stress. Thus TabZIP (Traes_7AL_25850F96F.1) can serve as a candidate gene for improving heat as well as other abiotic stress tolerance and can be helpful in enhancing the crop productivity under stress conditions.

Genome-wide identification and characterization of gene family for RWP-RK transcription factors in wheat (Triticum aestivum L.).

RWP-RKs represent a small family of transcription factors (TFs) that are unique to plants and function particularly under conditions of nitrogen starvation. These RWP-RKs have been classified in two sub-families, NLPs (NIN-like proteins) and RKDs (RWP-RK domain proteins). NLPs regulate tissue-specific expression of genes involved in nitrogen use efficiency (NUE) and RKDs regulate expression of genes involved in gametogenesis/embryogenesis. During the present study, using in silico approach, 37 wheat RWP-RK genes were identified, which included 18 TaNLPs (2865 to 7340 bp with 4/5 exons), distributed on 15 chromosomes from 5 homoeologous groups (with two genes each on 4B,4D and 5A) and 19 TaRKDs (1064 to 5768 bp with 1 to 6 exons) distributed on 12 chromosomes from 4 homoeologous groups (except groups 1, 4 and 5); 2–3 splice variants were also available in 9 of the 37 genes. Sixteen (16) of these genes also carried 24 SSRs (simple sequence repeats), while 11 genes had targets for 13 different miRNAs. At the protein level, MD simulation analysis suggested their interaction with nitrate-ions. Significant differences were observed in the expression of only two (TaNLP1 and TaNLP2) of the nine representative genes that were used for in silico expression analysis under varying levels of N at post-anthesis stage (data for other genes was not available for in silico expression analysis). Differences in expression were also observed during qRT-PCR, when expression of four representative genes (TaNLP2, TaNLP7, TaRKD6 and TaRKD9) was examined in roots and shoots of seedlings (under different conditions of N supply) in two contrasting genotypes which differed in NUE (C306 with low NUE and HUW468 with high NUE). These four genes for qRT-PCR were selected on the basis of previous literature, level of homology and the level of expression (in silico study). In particular, the TaNLP7 gene showed significant up-regulation in the roots and shoots of HUW468 (with higher NUE) during N-starvation; this gene has already been characterized in Arabidopsis and tobacco, and is known to be involved in nitrate-signal transduction pathway.

Genome-Wide Identification and Expression Profiling of the TCP Family Genes in Spike and Grain Development of Wheat (Triticum aestivum L.).

The TCP family genes are plant-specific transcription factors and play important roles in plant development. TCPs have been evolutionarily and functionally studied in several plants. Although common wheat (Triticum aestivum L.) is a major staple crop worldwide, no systematic analysis of TCPs in this important crop has been conducted. Here, we performed a genome-wide survey in wheat and found 66 TCP genes that belonged to 22 homoeologous groups. We then mapped these genes on wheat chromosomes and found that several TCP genes were duplicated in wheat including the ortholog of the maize TEOSINTE BRANCHED 1. Expression study using both RT-PCR and in situ hybridization assay showed that most wheat TCP genes were expressed throughout development of young spike and immature seed. Cis-acting element survey along promoter regions suggests that subfunctionalization may have occurred for homoeologous genes. Moreover, protein–protein interaction experiments of three TCP proteins showed that they can form either homodimers or heterodimers. Finally, we characterized two TaTCP9 mutants from tetraploid wheat. Each of these two mutant lines contained a premature stop codon in the A subgenome homoeolog that was dominantly expressed over the B subgenome homoeolog. We observed that mutation caused increased spike and grain lengths. Together, our analysis of the wheat TCP gene family provides a start point for further functional study of these important transcription factors in wheat.