Institutional Publications
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Item MEDIATOR SUBUNIT17 is required for transcriptional optimization of root system architecture in Arabidopsis(Oxford University Press, 2023) Agrawal, Rekha; Singh, Amrita; Giri, Jitender; Magyar, Zoltan; Thakur, Jitendra K.Sucrose and auxin are well-known determinants of root system architecture (RSA). However, the factors that connect the signaling pathways evoked by these two critical factors during root development are poorly understood. In this study, we report the role of MEDIATOR SUBUNIT17 (MED17) in RSA and its involvement in the transcriptional integration of sugar and auxin signaling pathways in Arabidopsis (Arabidopsis thaliana). Sucrose regulates root meristem activation through the TARGET OF RAPAMYCIN-E2 PROMOTER BINDING FACTOR A TOR-E2FA pathway, and auxin regulates lateral root (LR) development through AUXIN RESPONSE FACTOR-LATERAL ORGAN BOUNDARIES DOMAIN ARF-LBDs. Both sucrose and auxin play a vital role during primary and LR development. However, there is no clarity on how sucrose is involved in the ARF-dependent regulation of auxin-responsive genes. This study establishes MED17 as a nodal point to connect sucrose and auxin signaling. Transcription of MED17 was induced by sucrose in an E2FA/B-dependent manner. Moreover, E2FA/B interacted with MED17, which can aid in the recruitment of the Mediator complex on the target promoters. Interestingly, E2FA/B and MED17 also occupied the promoter of ARF7, but not ARF19, leading to ARF7 expression, which then activates auxin signaling and thus initiates LR development. MED17 also activated cell division in the root meristem by occupying the promoters of cell-cycle genes, thus regulating their transcription. Thus, MED17 plays an important role in relaying the transcriptional signal from sucrose to auxin-responsive and cell-cycle genes to regulate primary and lateral root development, highlighting the role of the Mediator as the transcriptional processor for optimal root system architecture in Arabidopsis.Item MEDIATOR SUBUNIT17 integrates jasmonate and auxin signaling pathways to regulate thermomorphogenesis(Oxford University Press, 2022) Agrawal, Rekha; Sharma, Mohan; Dwivedi, Nidhi; Maji, Sourobh; Thakur, Pallabi; Junaid, Alim; Fajkus, Jiří; Laxmi, Ashverya; Thakur, Jitendra K.Plant adjustment to environmental changes involves complex crosstalk between extrinsic and intrinsic cues. In the past two decades, extensive research has elucidated the key roles of PHYTOCHROME INTERACTING FACTOR4 (PIF4) and the phytohormone auxin in thermomorphogenesis. In this study, we identified a previously unexplored role of jasmonate (JA) signaling components, the Mediator complex, and their integration with auxin signaling during thermomorphogenesis in Arabidopsis (Arabidopsis thaliana). Warm temperature induces expression of JA signaling genes including MYC2, but, surprisingly, this transcriptional activation is not JA dependent. Warm temperature also promotes accumulation of the JA-signaling receptor CORONATINE INSENSITIVE1 (COI1) and degradation of the JA-signaling repressor JASMONATE-ZIM-DOMAIN PROTEIN9 (JAZ9), which probably leads to de-repression of MYC2, enabling it to contribute to the expression of MEDIATOR SUBUNIT17 (MED17). In response to warm temperature, MED17 occupies the promoters of thermosensory genes including PIF4, YUCCA8 (YUC8), INDOLE-3-ACETIC ACID INDUCIBLE19 (IAA19), and IAA29. Moreover, MED17 facilitates enrichment of H3K4me3 on the promoters of PIF4, YUC8, IAA19, and IAA29 genes. Interestingly, both occupancy of MED17 and enrichment of H3K4me3 on these thermomorphogenesis-related promoters are dependent on PIF4 (or PIFs). Altered accumulation of COI1 under warm temperature in the med17 mutant suggests the possibility of a feedback mechanism. Overall, this study reveals the role of the Mediator complex as an integrator of JA and auxin signaling pathways during thermomorphogenesis.Item Kinase module of mediator complex: important signalling processor for development and survival of plants(Oxford University Press, 2021) Agrawal, Rekha; Jiří, Fajkus; Thakur, Jitendra K.Mediator, a multisubunit protein complex is a signal processor that conveys regulatory information from transcription factors (TFs) to RNA Polymerase II (RNA Pol II) and therefore, plays an important role in the regulation of gene expression. This mega Dalton complex comprises four modules namely Head, Middle, Tail, and Kinase. The first three modules form the core part of the complex whereas association of the Kinase module is facultative. Being able to tweak the function of Mediator, the Kinase module has established itself as a major transcriptional regulator of numerous developmental and biochemical processes. The Kinase module consists of MED12, MED13, CycC, and kinase CDK8. Upon association with Mediator, the Kinase module can alter its structure and function dramatically. In the last decade, it has been established that the Kinase module is very important for plant growth and development and the fight against biotic and abiotic challenges. However, there is no comprehensive review discussing these findings in detail and depth. In this review, we have surveyed the regulation of Kinase module subunits and highlighted the multitude of their functions in plants. Coordination between the subunits to process different signals for optimum plant growth and development is also discussed.Item Interaction map of Arabidopsis mediator complex expounding its topology(Oxford University Press, 2019) Maji, Sourobh; Dahiya, Pradeep; Waseem, Mohd; Dwivedi, Nidhi; Bhat, Divya S.; Dar, Tanvir H.; Thakur, Jitendra K.Understanding of mechanistic details of Mediator functioning in plants is impeded as the knowledge of subunit organization and structure is lacking. In this study, an interaction map of Arabidopsis Mediator complex was analyzed to understand the arrangement of the subunits in the core part of the complex. Combining this interaction map with homologybased modeling, probable structural topology of core part of the Arabidopsis Mediator complex was deduced. Though the overall topology of the complex was similar to that of yeast, several differences were observed. Many interactions discovered in this study are not yet reported in other systems. AtMed14 and AtMed17 emerged as the key component providing important scaffold for the whole complex. AtMed6 and AtMed10 were found to be important for linking head with middle and middle with tail, respectively. Some Mediator subunits were found to form homodimers and some were found to possess transactivation property. Subcellular localization suggested that many of the Mediator subunits might have functions beyond the process of transcription. Overall, this study reveals role of individual subunits in the organization of the core complex, which can be an important resource for understanding the molecular mechanism of functioning of Mediator complex and its subunits in plants.Item Role of plant mediator complex in stress response(Springer, 2015) Samanta, Subhasis; Thakur, Jitendra K.Class II gene loci of eukaryotes are transcribed by RNA Polymerase II, which functions in coordination with several other proteins like transcription factors, general transcription factors, and cofactors. Recently, Mediator complex, a multi-subunit, megadalton size protein complex has gained lots of attention as an important component of RNA pol II transcriptional machinery because of its essentiality in the regulation of most of the class II genes. Like yeast and other metazoans, plants also possess the Mediator complex across the kingdom, and its isolation and subunit analyses have been reported from the model plant, Arabidopsis. Recent times have experienced a flurry of scientific papers containing the functional information of individual Mediator subunits in plants, although many were reported earlier without consideration of their association with the Mediator complex. Among its diverse functional aspects, several reports have established the Mediator complex as an important integrative hub of different biotic and abiotic stress signaling pathways, which have been discussed in this chapter from the functional genomics perspectives. Although reports are emerging in support of its inclusion as a component of the basic transcriptional machinery, the gene selective roles of the individual Mediator subunits are proven and indisputably accepted.Item Importance of mediator complex in the regulation and integration of diverse signaling pathways in plants(Frontiers Media S.A., 2015) Samanta, Subhasis; Thakur, Jitendra K.Basic transcriptional machinery in eukaryotes is assisted by a number of cofactors, which either increase or decrease the rate of transcription. Mediator complex is one such cofactor, and recently has drawn a lot of interest because of its integrative power to converge different signaling pathways before channeling the transcription instructions to the RNA polymerase II machinery. Like yeast and metazoans, plants do possess the Mediator complex across the kingdom, and its isolation and subunit analyses have been reported from the model plant, Arabidopsis. Genetic, and molecular analyses have unraveled important regulatory roles of Mediator subunits at every stage of plant life cycle starting from flowering to embryo and organ development, to even size determination. It also contributes immensely to the survival of plants against different environmental vagaries by the timely activation of its resistance mechanisms. Here, we have provided an overview of plant Mediator complex starting from its discovery to regulation of stoichiometry of its subunits. We have also reviewed involvement of different Mediator subunits in different processes and pathways including defense response pathways evoked by diverse biotic cues. Wherever possible, attempts have been made to provide mechanistic insight of Mediator's involvement in these processes.Item Sequence and expression analyses of KIX domain proteins suggest their importance in seed development and determination of seed size in rice, and genome stability in Arabidopsis(Springer, 2013) Thakur, Jitendra K.; Agarwal, Pinky; Parida, Swarup K.; Bajaj, Deepak; Pasrija, RichaThe KIX domain, which mediates protein-protein interactions, was first discovered as a motif in the large multidomain transcriptional activator histone acetyltransferase p300/CBP. Later, the domain was also found in Mediator subunit MED15, where it interacts with many transcription factors. In both proteins, the KIX domain is a target of activation domains of diverse transcription activators. It was found to be an essential component of several specific gene-activation pathways in fungi and metazoans. Not much is known about KIX domain proteins in plants. This study aims to characterize all the KIX domain proteins encoded by the genomes of Arabidopsis and rice. All identified KIX domain proteins are presented, together with their chromosomal locations, phylogenetic analysis, expression and SNP analyses. KIX domains were found not only in p300/CBP- and MED15-like plant proteins, but also in F-box proteins in rice and DNA helicase in Arabidopsis, suggesting roles of KIX domains in ubiquitin-mediated proteasomal degradation and genome stability. Expression analysis revealed overlapping expression of OsKIX_3, OsKIX_5 and OsKIX_7 in different stages of rice seeds development. Moreover, an association analysis of 136 in silico mined SNP loci in 23 different rice genotypes with grain-length information identified three non-synonymous SNP loci in these three rice genes showing strong association with long- and short-grain differentiation. Interestingly, these SNPs were located within KIX domain encoding sequences. Overall, this study lays a foundation for functional analysis of KIX domain proteins in plants.Item Tissue specific expression profile of mediator genes in Arabidopsis(Landes Bioscience (Taylor & Francis Grp), 2013) Pasrija, Richa; Thakur, Jitendra K.Mediator is a gigantic multiprotein complex required for transcription of almost all the protein coding genes. In this report, we have analyzed the transcript abundance of 31 Med genes in different tissues of Arabidopsis. Our analysis revealed the tissue specific differential expression profile of many Med subunit genes suggesting they might be contributing in the formation, maturation or function of that specific tissue. Moreover, we also found increase or decrease in the expression level of several Med subunits during the same duration of specific processes (for example flowering) indicating probable enrichment of a particular arrangement of selected subunits during that process. Thus, this study suggests that not only specific Med subunits have functional relevance in specific processes, but specific arrangements of Med subunits might also play significant role in some processes in Arabidopsis or other plants.