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The WRKY transcription factor family in Brachypodium distachyon
Prateek Tripathi, Roel C Rabara, Tanner J Langum, Ashley K Boken, Deena L Rushton, Darius D Boomsma, Charles I Rinerson, Jennifer Rabara, R Neil Reese, Xianfeng Chen, Jai S Rohila, Paul J Rushton
BMC Genomics , 2012, DOI: 10.1186/1471-2164-13-270
Abstract: We have identified and manually curated the WRKY transcription factor family from Brachypodium using a pipeline designed to identify all potential WRKY genes. 86 WRKY transcription factors were found, a total higher than all other current databases. We therefore propose that our numbering system (BdWRKY1-BdWRKY86) becomes the standard nomenclature. In the JGI v1.0 assembly of Brachypodium with the MIPS/JGI v1.0 annotation, nine of the transcription factors have no gene model and eleven gene models are probably incorrectly predicted. In total, twenty WRKY transcription factors (23.3%) do not appear to have accurate gene models. To facilitate use of our data, we have produced The Database of Brachypodium distachyon WRKY Transcription Factors. Each WRKY transcription factor has a gene page that includes predicted protein domains from MEME analyses. These conserved protein domains reflect possible input and output domains in signaling. The database also contains a BLAST search function where a large dataset of WRKY transcription factors, published genes, and an extensive set of wheat ESTs can be searched. We also produced a phylogram containing the WRKY transcription factor families from Brachypodium, rice, Arabidopsis, soybean, and Physcomitrella patens, together with published WRKY transcription factors from wheat. This phylogenetic tree provides evidence for orthologues, co-orthologues, and paralogues of Brachypodium WRKY transcription factors.The description of the WRKY transcription factor family in Brachypodium that we report here provides a framework for functional genomics studies in an important model system. Our database is a resource for both Brachypodium and wheat studies and ultimately projects aimed at improving wheat through manipulation of WRKY transcription factors.
Phylogenomic analysis of the cystatin superfamily in eukaryotes and prokaryotes
Du?an Kordi?, Vito Turk
BMC Evolutionary Biology , 2009, DOI: 10.1186/1471-2148-9-266
Abstract: We have identified in silico the full complement of the cystatin superfamily in more than 2100 prokaryotic and eukaryotic genomes. The analysis of numerous eukaryotic genomes has provided strong evidence for the emergence of this superfamily in the ancestor of eukaryotes. The progenitor of this superfamily was most probably intracellular and lacked a signal peptide and disulfide bridges, much like the extant Giardia cystatin. A primordial gene duplication produced two ancestral eukaryotic lineages, cystatins and stefins. While stefins remain encoded by a single or a small number of genes throughout the eukaryotes, the cystatins have undergone a more complex and dynamic evolution through numerous gene and domain duplications. In the cystatin superfamily we discovered twenty vertebrate-specific and three angiosperm-specific orthologous families, indicating that functional diversification has occurred only in multicellular eukaryotes. In vertebrate orthologous families, the prevailing trends were loss of the ancestral inhibitory activity and acquisition of novel functions in innate immunity. Bacterial cystatins and stefins may be emergency inhibitors that enable survival of bacteria in the host, defending them from the host's proteolytic activity.This study challenges the current view on the classification, origin and evolution of the cystatin superfamily and provides valuable insights into their functional diversification. The findings of this comprehensive study provide guides for future structural and evolutionary studies of the cystatin superfamily as well as of other protease inhibitors and proteases.The cystatin superfamily consists of a large group of cystatin domain-containing proteins, most of which are reversible and tight-binding inhibitors of the papain (C1) and legumain (C13) families of cysteine proteases [1-4]. On the basis of sequence similarity, the presence or lack of disulfide bonds, and physiological localization, this superfamily has been divided i
Roles of arabidopsis WRKY18, WRKY40 and WRKY60 transcription factors in plant responses to abscisic acid and abiotic stress
Han Chen, Zhibing Lai, Junwei Shi, Yong Xiao, Zhixiang Chen, Xinping Xu
BMC Plant Biology , 2010, DOI: 10.1186/1471-2229-10-281
Abstract: We report that the three WRKYs are involved in plant responses to abscisic acid (ABA) and abiotic stress. Through analysis of single, double, and triple mutants and overexpression lines for the WRKY genes, we have shown that WRKY18 and WRKY60 have a positive effect on plant ABA sensitivity for inhibition of seed germination and root growth. The same two WRKY genes also enhance plant sensitivity to salt and osmotic stress. WRKY40, on the other hand, antagonizes WRKY18 and WRKY60 in the effect on plant sensitivity to ABA and abiotic stress in germination and growth assays. Both WRKY18 and WRKY40 are rapidly induced by ABA, while induction of WRKY60 by ABA is delayed. ABA-inducible expression of WRKY60 is almost completely abolished in the wrky18 and wrky40 mutants. WRKY18 and WRKY40 recognize a cluster of W-box sequences in the WRKY60 promoter and activate WRKY60 expression in protoplasts. Thus, WRKY60 might be a direct target gene of WRKY18 and WRKY40 in ABA signaling. Using a stable transgenic reporter/effector system, we have shown that both WRKY18 and WRKY60 act as weak transcriptional activators while WRKY40 is a transcriptional repressor in plant cells.We propose that the three related WRKY transcription factors form a highly interacting regulatory network that modulates gene expression in both plant defense and stress responses by acting as either transcription activator or repressor.Plants are constantly exposed to a variety of biotic and abiotic stresses and have evolved intricate mechanisms to sense and respond to the adverse conditions. Phytohormones such as salicylic acid (SA), ethylene (ET), jasmonic acid (JA) and abscisic acid (ABA) play important roles in the regulation of plant responses to the adverse environmental conditions. In Arabidopsis, mutants deficient in SA biosynthesis (e.g. sid2) or signalling (e.g. npr1) exhibit enhanced susceptibility to biotrophic pathogens, which parasitize on plant living tissue [1,2]. ET- and JA-mediated signaling pat
Roles of Arabidopsis WRKY3 and WRKY4 Transcription Factors in Plant Responses to Pathogens
Zhibing Lai, KM Vinod, Zuyu Zheng, Baofang Fan, Zhixiang Chen
BMC Plant Biology , 2008, DOI: 10.1186/1471-2229-8-68
Abstract: Both WRKY3 and WRKY4 are nuclear-localized and specifically recognize the TTGACC W-box sequences in vitro. Expression of WRKY3 and WRKY4 was induced rapidly by stress conditions generated by liquid infiltration or spraying. Stress-induced expression of WRKY4 was further elevated by pathogen infection and SA treatment. To determine directly their role in plant disease resistance, we have isolated T-DNA insertion mutants and generated transgenic overexpression lines for WRKY3 and WRKY4. Both the loss-of-function mutants and transgenic overexpression lines were examined for responses to the biotrophic bacterial pathogen Pseudomonas syringae and the necrotrophic fungal pathogen Botrytis cinerea. The wrky3 and wrky4 single and double mutants exhibited more severe disease symptoms and support higher fungal growth than wild-type plants after Botrytis infection. Although disruption of WRKY3 and WRKY4 did not have a major effect on plant response to P. syringae, overexpression of WRKY4 greatly enhanced plant susceptibility to the bacterial pathogen and suppressed pathogen-induced PR1 gene expression.The nuclear localization and sequence-specific DNA-binding activity support that WRKY3 and WRKY4 function as transcription factors. Functional analysis based on T-DNA insertion mutants and transgenic overexpression lines indicates that WRKY3 and WRKY4 have a positive role in plant resistance to necrotrophic pathogens and WRKY4 has a negative effect on plant resistance to biotrophic pathogens.Upon pathogen infection, pathogen-associated molecular patterns (PAMPs) such as bacterial flagellin and lipopolysaccharides are recognized by plant receptors to activate PAMP-triggered immunity through a mitogen-activated protein kinase signaling cascade [1]. Gram-negative bacterial pathogens such as Pseudomonas syringae can deliver effector proteins to plant cells to interfere PAMP-triggered resistance to promote pathogen virulence. As a result, the remaining basal defense is usually insuffi
Expression Analysis of WRKY Transcription Factor Genes in Response to Abiotic Stresses in Horsegram (Macrotyloma uniflorum (Lam.) Verdc.)  [PDF]
Kurnool Kiranmai, Lokanadha Rao Gunupuru, Ambekar Nareshkumar, Vennapusa Amaranatha Reddy, Uppala Lokesh, Merum Pandurangaiah, Boya Venkatesh, Tanguturi Venkata Kirankumar, Chinta Sudhakar
American Journal of Molecular Biology (AJMB) , 2016, DOI: 10.4236/ajmb.2016.64013
Abstract: Drought and salt stress are two major environmental constraints that limit the productivity of agriculture crops worldwide. WRKY transcription factors are the plant-specific transcription factors that regulate several developmental events and stress responses in plants. The WRKY domain is defined by a 60-amino acid conserved sequence named WRKYGQK at N-terminal and a Zinc Finger-like motif at the C-terminal. WRKY genes are known to respond several stresses which may act as negative or positive regulators. The function of most of the WRKY transcription factors from non-model plants remains poorly understood. This investigation shows the expression levels of eight WRKY transcription factor genes from horsegram plant under drought and salt stress conditions. The increase in mRNA transcript levels of WRKY transcription factor genes was found to be high in drought stressed plants compared to salt-stressed plants. The levels of MDA which indicates the lipid peroxidation were less in drought stress. More ROS is produced in salt stress conditions compared to drought. The results show that the expression of WRKY transcription factors in drought stress conditions is reducing the adverse effect of stress on plants. These results also suggest that, during abiotic stress conditions such as drought and salt stress, WRKY transcription factors are regulated at the transcription level.
WRKY Transcription Factors Involved in Activation of SA Biosynthesis Genes
Marcel C van Verk, John F Bol, Huub JM Linthorst
BMC Plant Biology , 2011, DOI: 10.1186/1471-2229-11-89
Abstract: Expression studies with ICS1 promoter::β-glucuronidase (GUS) genes in Arabidopsis thaliana protoplasts cotransfected with 35S::WRKY28 showed that over expression of WRKY28 resulted in a strong increase in GUS expression. Moreover, qRT-PCR analyses indicated that the endogenous ICS1 and PBS3 genes were highly expressed in protoplasts overexpressing WRKY28 or WRKY46, respectively. Electrophoretic mobility shift assays indentified potential WRKY28 binding sites in the ICS1 promoter, positioned -445 and -460 base pairs upstream of the transcription start site. Mutation of these sites in protoplast transactivation assays showed that these binding sites are functionally important for activation of the ICS1 promoter. Chromatin immunoprecipitation assays with haemagglutinin-epitope-tagged WRKY28 showed that the region of the ICS1 promoter containing the binding sites at -445 and -460 was highly enriched in the immunoprecipitated DNA.The results obtained here confirm results from our multiple microarray co-expression analyses indicating that WRKY28 and WRKY46 are transcriptional activators of ICS1 and PBS3, respectively, and support this in silico screening as a powerful tool for identifying new components of stress signaling pathways.Because of their sessile nature, plants have evolved very sophisticated mechanisms to actively cope with different sorts of stresses. The various defense mechanisms are controlled by signaling molecules like salicylic acid (SA), jasmonic acid (JA), and ethylene, or by combinations of these signal compounds. SA accumulates locally in infected leaves, as well as in non-infected systemic leaves after infection with biotrophic pathogens and mediates the induced expression of defense genes, resulting in an enhanced state of defense known as systemic acquired resistance (SAR) [1-5]. SAR is a long-lasting broad-spectrum resistance against a variety of pathogenic fungi, bacteria and viruses [6,7]. Also exogenous application of SA results in induced exp
Phosphorylation of a WRKY Transcription Factor by MAPKs Is Required for Pollen Development and Function in Arabidopsis  [PDF]
Yuefeng Guan,Xiangzong Meng,Reshma Khanna,Erica LaMontagne,Yidong Liu,Shuqun Zhang
PLOS Genetics , 2014, DOI: doi/10.1371/journal.pgen.1004384
Abstract: Plant male gametogenesis involves complex and dynamic changes in gene expression. At present, little is known about the transcription factors involved in this process and how their activities are regulated. Here, we show that a pollen-specific transcription factor, WRKY34, and its close homolog, WRKY2, are required for male gametogenesis in Arabidopsis thaliana. When overexpressed using LAT52, a strong pollen-specific promoter, epitope-tagged WRKY34 is temporally phosphorylated by MPK3 and MPK6, two mitogen-activated protein kinases (MAPKs, or MPKs), at early stages in pollen development. During pollen maturation, WRKY34 is dephosphorylated and degraded. Native promoter-driven WRKY34-YFP fusion also follows the same expression pattern at the protein level. WRKY34 functions redundantly with WRKY2 in pollen development, germination, and pollen tube growth. Loss of MPK3/MPK6 phosphorylation sites in WRKY34 compromises the function of WRKY34 in vivo. Epistasis interaction analysis confirmed that MPK6 belongs to the same genetic pathway of WRKY34 and WRKY2. Our study demonstrates the importance of temporal post-translational regulation of WRKY transcription factors in the control of developmental phase transitions in plants.
The Beet Cyst Nematode Heterodera schachtii Modulates the Expression of WRKY Transcription Factors in Syncytia to Favour Its Development in Arabidopsis Roots  [PDF]
Muhammad Amjad Ali, Krzysztof Wieczorek, David P. Kreil, Holger Bohlmann
PLOS ONE , 2014, DOI: 10.1371/journal.pone.0102360
Abstract: Cyst nematodes invade the roots of their host plants as second stage juveniles and induce a syncytium which is the only source of nutrients throughout their life. A recent transcriptome analysis of syncytia induced by the beet cyst nematode Heterodera schachtii in Arabidopsis roots has shown that thousands of genes are up-regulated or down-regulated in syncytia as compared to root segments from uninfected plants. Among the down-regulated genes are many which code for WRKY transcription factors. Arabidopsis contains 66 WRKY genes with 59 represented by the ATH1 GeneChip. Of these, 28 were significantly down-regulated and 6 up-regulated in syncytia as compared to control root segments. We have studied here the down-regulated genes WRKY6, WRKY11, WRKY17 and WRKY33 in detail. We confirmed the down-regulation in syncytia with promoter::GUS lines. Using various overexpression lines and mutants it was shown that the down-regulation of these WRKY genes is important for nematode development, probably through interfering with plant defense reactions. In case of WRKY33, this might involve the production of the phytoalexin camalexin.
Genome-wide analysis of WRKY gene family in Cucumis sativus
Jian Ling, Weijie Jiang, Ying Zhang, Hongjun Yu, Zhenchuan Mao, Xingfang Gu, Sanwen Huang, Bingyan Xie
BMC Genomics , 2011, DOI: 10.1186/1471-2164-12-471
Abstract: We identified a total of 55 WRKY genes in the cucumber genome. According to structural features of their encoded proteins, the cucumber WRKY (CsWRKY) genes were classified into three groups (group 1-3). Analysis of expression profiles of CsWRKY genes indicated that 48 WRKY genes display differential expression either in their transcript abundance or in their expression patterns under normal growth conditions, and 23 WRKY genes were differentially expressed in response to at least one abiotic stresses (cold, drought or salinity). The expression profile of stress-inducible CsWRKY genes were correlated with those of their putative Arabidopsis WRKY (AtWRKY) orthologs, except for the group 3 WRKY genes. Interestingly, duplicated group 3 AtWRKY genes appear to have been under positive selection pressure during evolution. In contrast, there was no evidence of recent gene duplication or positive selection pressure among CsWRKY group 3 genes, which may have led to the expressional divergence of group 3 orthologs.Fifty-five WRKY genes were identified in cucumber and the structure of their encoded proteins, their expression, and their evolution were examined. Considering that there has been extensive expansion of group 3 WRKY genes in angiosperms, the occurrence of different evolutionary events could explain the functional divergence of these genes.Transcription factors exhibit sequence-specific DNA-binding and are capable of activating or repressing transcription of downstream target genes. In plants, WRKY proteins constitute a large family of transcription factors that are involved in various physiological processes. Proteins in this family contain at least one highly conserved signature domain of about 60 amino acid residues, which includes the conserved WRKYGQK sequence followed by a zinc finger motif, located in the C-terminal region [1]. The WRKY domain facilitates binding of the proteins to the W box or the SURE (sugar-responsive cis-element) in the promoter regions of
Massive expansion of the calpain gene family in unicellular eukaryotes  [cached]
Zhao Sen,Liang Zhe,Demko Viktor,Wilson Robert
BMC Evolutionary Biology , 2012, DOI: 10.1186/1471-2148-12-193
Abstract: Background Calpains are Ca2+-dependent cysteine proteases that participate in a range of crucial cellular processes. Dysfunction of these enzymes may cause, for instance, life-threatening diseases in humans, the loss of sex determination in nematodes and embryo lethality in plants. Although the calpain family is well characterized in animal and plant model organisms, there is a great lack of knowledge about these genes in unicellular eukaryote species (i.e. protists). Here, we study the distribution and evolution of calpain genes in a wide range of eukaryote genomes from major branches in the tree of life. Results Our investigations reveal 24 types of protein domains that are combined with the calpain-specific catalytic domain CysPc. In total we identify 41 different calpain domain architectures, 28 of these domain combinations have not been previously described. Based on our phylogenetic inferences, we propose that at least four calpain variants were established in the early evolution of eukaryotes, most likely before the radiation of all the major supergroups of eukaryotes. Many domains associated with eukaryotic calpain genes can be found among eubacteria or archaebacteria but never in combination with the CysPc domain. Conclusions The analyses presented here show that ancient modules present in prokaryotes, and a few de novo eukaryote domains, have been assembled into many novel domain combinations along the evolutionary history of eukaryotes. Some of the new calpain genes show a narrow distribution in a few branches in the tree of life, likely representing lineage-specific innovations. Hence, the functionally important classical calpain genes found among humans and vertebrates make up only a tiny fraction of the calpain family. In fact, a massive expansion of the calpain family occurred by domain shuffling among unicellular eukaryotes and contributed to a wealth of functionally different genes.
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