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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 The versatile role of glucose signalling in regulating growth, development and stress responses in plants(Springer Nature Publishing AG, 2020) Saksena, Harshita B.; Sharma, Manvi; Singh, Dhriti; Laxmi, AshveryaSugars as an energy source and a signalling molecule are indispensible for growth, development and stress responses in plants. Among sugars, glucose (Glc) has been identified to perform a key role in regulating various cellular and developmental processes in plants. This comprehensive review highlights the crosstalk of glucose signalling with auxin, cytokinin and brassinosteroid signalling in controlling several aspects of plant development including root growth deviation, root waving and coiling, lateral root emergence, hypocotyl elongation, etc. through a Hexokinase 1 dependent and independent pathway in Arabidopsis. It also provides an insight on the role of Glc-mediated target of rapamycin signalling in modulating various abiotic stresses like nutrient deficiency stress, heat stress, oxidative stress etc. The review also discusses about sugar inducible FCS-Like Zinc Finger genes in modulating SNF1-related protein kinase 1 signalling to coordinate growth and stress responses in plants. Therefore, glucose signalling in concert with diverse signalling pathways and via regulating certain genes contributes to numerous cellular processes to control development and stress responses in plants.Item FCS-like zinc finger 6 and 10 repress SnRK1 signalling in Arabidopsis(John Wiley & Sons, 2018) Jamsheer, Muhammed K.; Sharma, Manvi; Singh, Dhriti; Mannully, Chanchal Thomas; Jindal, Sunita; Shukla, Brihaspati Narayan; Laxmi, AshveryaSNF1-Related Protein Kinase 1 (SnRK1) is a central regulator of plant growth during energy starvation. The FCS-Like Zinc finger (FLZ) proteins are recently identified adaptor proteins which facilitate the interaction of SnRK1 with other proteins. In this study, we identified that two starvation-induced FLZ genes, FLZ6 and FLZ10, work as repressors of SnRK1 signalling. The reduced expression of these genes resulted in an increase in the level of SnRK1α1, which is the major catalytic subunit of SnRK1. This resulted in a concomitant increase in the phosphorylated protein and SnRK1 activity in the flz6 and flz10 mutants. FLZ6 and FLZ10 specifically interact with SnRK1α subunits in the cytoplasmic foci which colocalized with the endoplasmic reticulum. In physiological assays, similar to SnRK1α1 overexpression line, flz mutants showed compromised growth. Further, the growth promotion in response to favourable growth conditions was found to be attenuated in the mutants. The enhanced SnRK1 activity in the mutants resulted in a reduction in the level of phosphorylated RIBOSOMAL S6 KINASE and the expression of E2Fa and its targets indicating that TARGET OF RAPAMYCIN-dependent promotion of protein synthesis and cell cycle progression is impaired. Taken together, this study uncovers a plant-specific modulation of SnRK1 signalling.Item Arabidopsis RSS1 mediates cross-talk between glucose and light signaling during hypocotyl elongation growth(Nature Publishing Group, 2017) Singh, Manjul; Gupta, Aditi; Singh, Dhriti; Khurana, Jitendra P.; Laxmi, AshveryaPlants possess exuberant plasticity that facilitates its ability to adapt and survive under challenging environmental conditions. The developmental plasticity largely depends upon cellular elongation which is governed by a complex network of environmental and phytohormonal signals. Here, we report role of glucose (Glc) and Glc-regulated factors in controlling elongation growth and shade response in Arabidopsis. Glc controls shade induced hypocotyl elongation in a dose dependent manner. We have identified a Glc repressed factor REGULATED BY SUGAR AND SHADE1 (RSS1) encoding for an atypical basic helix-loop-helix (bHLH) protein of unknown biological function that is required for normal Glc actions. Phenotype analysis of mutant and overexpression lines suggested RSS1 to be a negative regulator of elongation growth. RSS1 affects overall auxin homeostasis. RSS1 interacts with the elongation growth-promoting proteins HOMOLOG OF BEE2 INTERACTING WITH IBH 1 (HBI1) and BR ENHANCED EXPRESSION2 (BEE2) and negatively affects the transcription of their downstream targets such as YUCs, INDOLE-3-ACETIC ACID INDUCIBLE (IAAs), LONG HYPOCOTYL IN FAR-RED1 (HFR1), HOMEOBOX PROTEIN 2 (ATHB2), XYLOGLUCAN ENDOTRANSGLUCOSYLASE/HYDROLASES (XTHs) and EXPANSINS. We propose, Glc signals might maintain optimal hypocotyl elongation under multiple signals such as light, shade and phytohormones through the central growth regulatory bHLH/HLH module.Item Striking the right chord: Signaling enigma during root gravitropism(Frontiers Media S.A., 2017) Singh, Manjul; Gupta, Aditi; Laxmi, AshveryaPlants being sessile can often be judged as passive acceptors of their environment. However, plants are actually even more active in responding to the factors from their surroundings. Plants do not have eyes, ears or vestibular system like animals, still they "know" which way is up and which way is down? This is facilitated by receptor molecules within plant which perceive changes in internal and external conditions such as light, touch, obstacles; and initiate signaling pathways that enable the plant to react. Plant responses that involve a definite and specific movement are called "tropic" responses. Perhaps the best known and studied tropisms are phototropism, i.e., response to light, and geotropism, i.e., response to gravity. A robust root system is vital for plant growth as it can provide physical anchorage to soil as well as absorb water, nutrients and essential minerals from soil efficiently. Gravitropic responses of both primary as well as lateral root thus become critical for plant growth and development. The molecular mechanisms of root gravitropism has been delved intensively, however, the mechanism behind how the potential energy of gravity stimulus converts into a biochemical signal in vascular plants is still unknown, due to which gravity sensing in plants still remains one of the most fascinating questions in molecular biology. Communications within plants occur through phytohormones and other chemical substances produced in plants which have a developmental or physiological effect on growth. Here, we review current knowledge of various intrinsic signaling mechanisms that modulate root gravitropism in order to point out the questions and emerging developments in plant directional growth responses. We are also discussing the roles of sugar signals and their interaction with phytohormone machinery, specifically in context of root directional responses.Item The interaction between glucose and cytokinin signaling in controlling Arabidopsis thaliana seedling root growth and development(Taylor & Francis Group, 2017) Kushwah, Sunita; Laxmi, AshveryaCytokinin (CK) and glucose (GLC) control a number of common responses in plants. There is an extensive overlap between CK and GLC signal transduction pathways in Arabidopsis. Physiologically, both GLC and CK could regulate root length in light. CK interacts with GLC via HXK1 dependent pathway for root length control. Wild-type (WT) roots cannot elongate in the GLC free medium while CK-receptor mutant ARABIDOPSIS HISTIDINE KINASE4 (ahk4) and type B ARR triple mutant ARABIDOPSIS RESPONSE REGULATOR1, 10,11 (arr1, 10,11) roots could elongate even in the absence of GLC as compared to the WT. The root hair initiation was also found defective in CK signaling mutants ahk4, arr1,10,11 and arr3,4,5,6,8,9 on increasing GLC concentration (up to 3%); and lesser number of root hairs were visible even at 5% GLC as compared to the WT. Out of 941 BAP regulated genes, 103 (11%) genes were involved in root growth and development. Out of these 103 genes, 60 (58%) genes were also regulated by GLC. GLC could regulate 5736 genes, which include 327 (6%) genes involved in root growth and development. Out of these 327 genes, 60 (18%) genes were also regulated by BAP. Both GLC and CK signaling cannot alter root length in light in auxin signaling mutant AUXIN RESPONSE3/INDOLE-3-ACETIC ACID17 (axr3/iaa17) suggesting that they may involve auxin signaling component as a nodal point. Therefore CK- and GLC- signaling are involved in controlling different aspects of root growth and development such as root length, with auxin signaling components working as downstream target.Item Expression of Arabidopsis FCS-like zinc finger genes is differentially regulated by sugars, cellular energy level, and abiotic stress(Frontiers Media S.A., 2015) K, Muhammed Jamsheer; Laxmi, AshveryaCellular energy status is an important regulator of plant growth, development, and stress mitigation. Environmental stresses ultimately lead to energy deficit in the cell which activates the SNF1-RELATED KINASE 1 (SnRK1) signaling cascade which eventually triggering a massive reprogramming of transcription to enable the plant to survive under low-energy conditions. The role of Arabidopsis thaliana FCS-Like Zinc finger (FLZ) gene family in energy and stress signaling is recently come to highlight after their interaction with kinase subunits of SnRK1 were identified. In a detailed expression analysis in different sugars, energy starvation, and replenishment series, we identified that the expression of most of the FLZ genes is differentially modulated by cellular energy level. It was found that FLZ gene family contains genes which are both positively and negatively regulated by energy deficit as well as energy-rich conditions. Genetic and pharmacological studies identified the role of HEXOKINASE 1- dependent and energy signaling pathways in the sugar-induced expression of FLZ genes. Further, these genes were also found to be highly responsive to different stresses as well as abscisic acid. In over-expression of kinase subunit of SnRK1, FLZ genes were found to be differentially regulated in accordance with their response toward energy fluctuation suggesting that these genes may work downstream to the established SnRK1 signaling under low-energy stress. Taken together, the present study provides a conceptual framework for further studies related to SnRK1-FLZ interaction in relation to sugar and energy signaling and stress responseItem Multiple interactions between glucose and brassinosteroid signal transduction pathways in Arabidopsis are uncovered by whole-genome transcriptional profiling(American Society of Plant Biologists, 2015) Gupta, Aditi; Singh, Manjul; Laxmi, AshveryaBrassinosteroid (BR) and glucose (Glc) regulate many common responses in plants. Here, we demonstrate that under etiolated growth conditions, extensive interdependence/overlap occurs between BR- and Glc-regulated gene expression as well as physiological responses. Glc could regulate the transcript level of 72% of BR-regulated genes at the whole-genome level, of which 58% of genes were affected synergistically and 42% of genes were regulated antagonistically. Presence of Glc along with BR in medium could affect BR induction/repression of 85% of BR-regulated genes. Glc could also regulate several genes involved in BR metabolism and signaling. Both BR and Glc coregulate a large number of genes involved in abiotic/biotic stress responses and growth and development. Physiologically, Glc and BR interact to regulate hypocotyl elongation growth of etiolated Arabidopsis (Arabidopsis thaliana) seedlings in a dose-dependent manner. Glc may interact with BR via a hexokinase1 (HXK1)-mediated pathway to regulate etiolated hypocotyl elongation. Brassinosteroid insensitive1 (BRI1) is epistatic to HXK1, as the Glc insensitive2bri1-6 double mutant displayed severe defects in hypocotyl elongation growth similar to its bri1-6 parent. Analysis of Glc and BR sensitivity in mutants defective in auxin response/signaling further suggested that Glc and BR signals may converge at S-phase kinase-associated protein1-Cullin-F-box-transport inhibitor response1/auxin-related f-box-auxin/indole-3-acetic acid-mediated auxin-signaling machinery to regulate etiolated hypocotyl elongation growth in Arabidopsis.Item Interaction between glucose and brassinosteroid during regulation of lateral root development in Arabidopsis thaliana(American Society of Plant Biologists, 2015) Gupta, Aditi; Singh, Manjul; Laxmi, AshveryaGlucose (Glc) plays a fundamental role in regulating lateral root (LR) development as well as LR emergence. In this study, we show that brassinosteroid (BR) signaling works downstream of Glc in controlling LR production/emergence in Arabidopsis (Arabidopsis thaliana) seedlings. Glc and BR can promote LR emergence at lower concentrations, while at higher concentrations, both have an inhibitory effect. The BR biosynthesis and perception mutants showed highly reduced numbers of emerged LRs at all the Glc concentrations tested. BR signaling works downstream of Glc signaling in regulating LR production, as in the glucose insensitive2-1brassinosteroid insensitive1 double mutant, Glc-induced LR production/emergence was severely reduced. Differential auxin distribution via the influx carriers AUXIN RESISTANT1/LIKE AUXIN RESISTANT1-3 and the efflux carrier PIN-FORMED2 plays a central role in controlling LR production in response to Glc and BR. Auxin signaling components AUXIN RESISTANT2,3 and SOLITARY ROOT act downstream of Glc and BR. AUXIN RESPONSE FACTOR7/19 work farther downstream and control LR production by regulating the expression of LATERAL ORGAN BOUNDARIES-DOMAIN29 and EXPANSIN17 genes. Increasing light flux could also mimic the Glc effect on LR production/emergence. However, increased light flux could not affect LR production in those BR and auxin signaling mutants that were defective for Glc-induced LR production. Altogether, our study suggests that, under natural environmental conditions, modulation of endogenous sugar levels can manipulate root architecture for optimized development by altering its nutrient/water uptake as well as its anchorage capacity.Item Glucose control of root growth direction in Arabidopsis thaliana(Oxford University Press, 2014) Singh, Manjul; Gupta, Aditi; Laxmi, AshveryaDirectional growth of roots is a complex process that is modulated by various environmental signals. This work shows that presence of glucose (Glc) in the medium also extensively modulated seedling root growth direction. Glc modulation of root growth direction was dramatically enhanced by simultaneous brassinosteroid (BR) application. Glc enhanced BR receptor BRASSINOSTEROID INSENSITIVE1 (BRI1) endocytosis from plasma membrane to early endosomes. Glc-induced root deviation was highly enhanced in a PP2A-defective mutant, roots curl in naphthyl phthalamic acid 1-1 (rcn1-1) suggesting that there is a role of phosphatase in Glc-induced root-growth deviation. RCN1, therefore, acted as a link between Glc and the BR-signalling pathway. Polar auxin transport worked further downstream to BR in controlling Glc-induced root deviation response. Glc also affected other root directional responses such as root waving and coiling leading to altered root architecture. High light intensity mimicked the Glc-induced changes in root architecture that were highly reduced in Glc-signalling mutants. Thus, under natural environmental conditions, changing light flux in the environment may lead to enhanced Glc production/response and is a way to manipulate root architecture for optimized development via integrating several extrinsic and intrinsic signalling cues.