Unfolded protein response (UPR) mediated under heat stress in plants

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REVIEW ARTICLE

Unfolded protein response (UPR) mediated under heat stress in plants M. K. Malini1 • V. S. Lekshmy2 • Madan Pal1 • Viswanathan Chinnusamy1 M. Nagaraj Kumar1



Received: 16 September 2020 / Accepted: 16 November 2020 Ó Indian Society for Plant Physiology 2020

Abstract Drastic fluctuation in temperature during crop growth challenges its yield to a greater extent, thus diminishing the agricultural productivity globally. At the cellular level, protein homeostasis is altered or disturbed directly or indirectly by the high temperature. Improper protein folding due to heat stress creates ER stress in the endoplasmic reticulum (ER). In order to balance the protein folding demand and its capacity and to maintain ER homeostasis, cells activate unfolded protein response (UPR), as a homeostatic mechanism. The transcription factors, bZip60 and bZip28/17 have been reported as the key modules of UPR for alleviating ER stress. Despite UPR mechanisms has been extensively investigated, the intervention of heat stress with ER homeostasis remains unclear. This review summarizes the key findings on this potential nexus. Additionally, the link between ER

Ramalingaswami Fellow-DBT. & M. Nagaraj Kumar [email protected] M. K. Malini [email protected] V. S. Lekshmy [email protected] Madan Pal [email protected] Viswanathan Chinnusamy [email protected] 1

Division of Plant Physiology, ICAR-Indian Agricultural Research Institute, New Delhi 110012, India

2

Department of Crop Physiology, University of Agricultural Sciences, GKVK, Bangalore, India

associated degradation and plant growth regulators during heat stress were also addressed. Keywords Basic leucine zipper (bZip)  Inositol-requiring enzyme 1(IRE1)  UPR (unfolded protein response)  ERAD (ER associated protein degradation)  Membraneassociated transcription factors (MTF)

Introduction Unpredictable climate anomalies accompanied with severe constraints on arable lands and available water enact serious threats to agriculture crops. The increased temperature prompts heat stress on plants and it seeks immediate attention for sustainable crop production due to its adverse impact on growth and development. Indeed, 1 °C raise in global mean temperature have drastically declined the yields of major crops (Zhao et al. 2017). Sensing elements are of utmost importance to understand the mechanisms of heat stress tolerance despite extensive reports on downstream genes and its regulation under heat stress were known. Plasma membrane (PM), a lipid-bilayer with intrinsic and transmembrane proteins, could be the site for the initiation of sensing external thermal fluctuations of the cell (Niu and Xiang 2018; Ruelland and Zachowski 2010). Heat stress affects the plasma membrane integrity through increased fluidity. The PM associated calcium channels sense this fluctuation and initiate the heat-shock responses (Demidchik et al. 2018; Ward et al. 2009) which are further amplified and orchestrated by the secondary transducers viz., histone sensors in