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      Drought Response in Wheat: Key Genes and Regulatory Mechanisms Controlling Root System Architecture and Transpiration Efficiency

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          Abstract

          Abiotic stresses such as, drought, heat, salinity, and flooding threaten global food security. Crop genetic improvement with increased resilience to abiotic stresses is a critical component of crop breeding strategies. Wheat is an important cereal crop and a staple food source globally. Enhanced drought tolerance in wheat is critical for sustainable food production and global food security. Recent advances in drought tolerance research have uncovered many key genes and transcription regulators governing morpho-physiological traits. Genes controlling root architecture and stomatal development play an important role in soil moisture extraction and its retention, and therefore have been targets of molecular breeding strategies for improving drought tolerance. In this systematic review, we have summarized evidence of beneficial contributions of root and stomatal traits to plant adaptation to drought stress. Specifically, we discuss a few key genes such as, DRO1 in rice and ERECTA in Arabidopsis and rice that were identified to be the enhancers of drought tolerance via regulation of root traits and transpiration efficiency. Additionally, we highlight several transcription factor families, such as, ERF (ethylene response factors), DREB (dehydration responsive element binding), ZFP (zinc finger proteins), WRKY, and MYB that were identified to be both positive and negative regulators of drought responses in wheat, rice, maize, and/or Arabidopsis. The overall aim of this review is to provide an overview of candidate genes that have been identified as regulators of drought response in plants. The lack of a reference genome sequence for wheat and non-transgenic approaches for manipulation of gene functions in wheat in the past had impeded high-resolution interrogation of functional elements, including genes and QTLs, and their application in cultivar improvement. The recent developments in wheat genomics and reverse genetics, including the availability of a gold-standard reference genome sequence and advent of genome editing technologies, are expected to aid in deciphering of the functional roles of genes and regulatory networks underlying adaptive phenological traits, and utilizing the outcomes of such studies in developing drought tolerant cultivars.

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          The role of stomata in sensing and driving environmental change.

          Stomata, the small pores on the surfaces of leaves and stalks, regulate the flow of gases in and out of leaves and thus plants as a whole. They adapt to local and global changes on all timescales from minutes to millennia. Recent data from diverse fields are establishing their central importance to plant physiology, evolution and global ecology. Stomatal morphology, distribution and behaviour respond to a spectrum of signals, from intracellular signalling to global climatic change. Such concerted adaptation results from a web of control systems, reminiscent of a 'scale-free' network, whose untangling requires integrated approaches beyond those currently used.
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            Control of root system architecture by DEEPER ROOTING 1 increases rice yield under drought conditions.

            The genetic improvement of drought resistance is essential for stable and adequate crop production in drought-prone areas. Here we demonstrate that alteration of root system architecture improves drought avoidance through the cloning and characterization of DEEPER ROOTING 1 (DRO1), a rice quantitative trait locus controlling root growth angle. DRO1 is negatively regulated by auxin and is involved in cell elongation in the root tip that causes asymmetric root growth and downward bending of the root in response to gravity. Higher expression of DRO1 increases the root growth angle, whereby roots grow in a more downward direction. Introducing DRO1 into a shallow-rooting rice cultivar by backcrossing enabled the resulting line to avoid drought by increasing deep rooting, which maintained high yield performance under drought conditions relative to the recipient cultivar. Our experiments suggest that control of root system architecture will contribute to drought avoidance in crops.
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              APETALA2/Ethylene Responsive Factor (AP2/ERF) transcription factors: mediators of stress responses and developmental programs.

              Transcription factors belonging to the APETALA2/Ethylene Responsive Factor (AP2/ERF) family are conservatively widespread in the plant kingdom. These regulatory proteins are involved in the control of primary and secondary metabolism, growth and developmental programs, as well as responses to environmental stimuli. Due to their plasticity and to the specificity of individual members of this family, AP2/ERF transcription factors represent valuable targets for genetic engineering and breeding of crops. In this review, we integrate the evidence collected from functional and structural studies to describe their different mechanisms of action and the regulatory pathways that affect their activity.
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                Author and article information

                Contributors
                Journal
                Front Chem
                Front Chem
                Front. Chem.
                Frontiers in Chemistry
                Frontiers Media S.A.
                2296-2646
                05 December 2017
                2017
                : 5
                : 106
                Affiliations
                [1] 1Canadian Wheat Improvement Flagship Program, National Research Council Canada (NRC-CNRC) , Saskatoon, SK, Canada
                [2] 2Saskatoon Research and Development Centre, Agriculture and Agri-Food Canada , Saskatoon, SK, Canada
                [3] 3Department of Global Agricultural Sciences, Graduate School of Agricultural and Life Sciences, The University of Tokyo , Tokyo, Japan
                [4] 4Institute of Crop Science (NICS), National Agriculture and Food Research Organization (NARO) , Tsukuba, Japan
                [5] 5Agrobioversity Research Area, International Center for Tropical Agriculture (CIAT) , Cali, Colombia
                Author notes

                Edited by: Raju Datla, National Research Council Canada (NRC-CNRC), Canada

                Reviewed by: Rupesh Kailasrao Deshmukh, Laval University, Canada; Noemi Tel Zur, Ben-Gurion University of the Negev, Beersheba, Israel

                *Correspondence: Sateesh Kagale Sateesh.Kagale@ 123456nrc-cnrc.gc.ca

                This article was submitted to Agricultural Biological Chemistry, a section of the journal Frontiers in Chemistry

                Article
                10.3389/fchem.2017.00106
                5723305
                29259968
                0680709f-3757-40c4-a178-7abcf0d556df
                Copyright © 2017 Kulkarni, Soolanayakanahally, Ogawa, Uga, Selvaraj and Kagale.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 31 August 2017
                : 07 November 2017
                Page count
                Figures: 1, Tables: 4, Equations: 0, References: 129, Pages: 13, Words: 11104
                Categories
                Chemistry
                Review

                wheat,drought,root traits,transpiration efficiency,transcriptional regulation,ear motif

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