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Phytohormones and Abiotic Stress Tolerance in Plants
Name: Phytohormones and Abiotic Stress Tolerance in Plants
Author: f. eyidogan
Pages: 311
Year: 2012
Language: English
File Size: 4.14 MB
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Editors Nafees A. Khan Rahat Nazar Noushina Iqbal Aligarh Muslim University Department of Botany Aligarh India [email protected] [email protected] [email protected] A. Anjum Centre for Environmental and Marine Stud Department of Chemistry Aveiro Portugal [email protected] ISBN 978 3 642 25828 2 e ISBN 978 3 642 25829 9 DOI 10.1007/978 3 642 25829 9 Springer Heidelberg Dordrecht London New York Library of Congress Control Number: 2012933369 #Springer Verlag Berlin Heidelberg 2012 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, speci cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on micro lms or in other ways, and storage in data banks. Duplication of this publication or partsthereofispermittedonlyundertheprovisionsoftheGermanCopyrightLawofSeptember9,1965,in itscurrentversion,andpermissionforusemustalwaysbeobtainedfromSpringer.Violationsare liableto prosecution under the German Copyright Law. The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply,evenintheabsenceofaspeci cstatement,thatsuchnamesareexemptfromtherelevantprotective laws and regulations and therefore free for general use. Printed on acid free paper Springer is part of Springer Science+Business Media (

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Preface Plants are exposed to rapid and various unpredicted disturbances in the environ ment resulting in stressful conditions. Abiotic stress is the negative impact of nonliving factors on the living organisms in a speci c environment and constitutes a major limitation to agricultural production. The adverse environmental conditions that plants encounter during their life cycle disturb metabolic reactions and ad versely affect growth and development at cellular and whole plant level. Under abiotic stress, plants integrate multiple external stress cues to bring about a coordi nated response and establish mechanism to mitigate the stress by triggering a cascade of events leading to enhanced tolerance. Responses to stress are complicat ed integrated circuits involving multiple pathways and speci c cellular compart ments, and the interaction of additional cofactors and/or signaling molecules coordinates a speci ed response to a given stimulus. Stress signal is rst perceived by the receptors present on the membrane of the plant cells. The signal information is then transduced downstream resulting in the activation of various stress responsive genes.The products of these stress genes ultimately lead tostresstolerance response or plant adaptation and help the plant to survive and surpass the unfavorable conditions. Abiotic stress conditions lead to production of signaling molecule(s) that induce the synthesis of several metabolites, including phytohormones for stress tolerance. Phytohormones are chemical compounds produced in one part and exert effect in another part and in uence physiological and biochemical processes. Phytohormones are critical for plant growth and development and play an important role in integrating various stress signals and controlling downstream stress responses and interact in coordination with each other for defense signal network ing to ne tune defense. The adaptive process of plants response imposed by abiotic stresses such as salt, cold, drought, and wounding is mainly controlled by the phytohormones. Stress conditions activate phytohormones signaling pathways that are thought to mediate adaptive responses at extremely low concentration. Thus, an understanding of the phytohormones homeostasis and signaling is essen tial for improving plant performance under optimal and stressful environments. v

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Traditionally ve major classes of plant hormones have been recognized: auxins, cytokinins, gibberellins, abscisic acid, and ethylene. Recently, other signaling molecules that play roles in plant metabolism and abiotic stress tolerance have also been identi ed, including brassinosteroids, jasmonic acid, salicylic acid, and nitric oxide. Besides, more active molecules are being found and new families of regulators are emerging such as polyamines, plant peptides, and karrikins. Several biological effects of phytohormones are induced by cooperation of more than one phytohormone. Substantial progress has been made in understanding individual aspects of phytohormones perception, signal transduction, homeostasis, or in u ence on gene expression. However, the physiological, biochemical, and molecular mechanisms induced by phytohormones through which plants integrate adaptive responses under abiotic stress are largely unknown. This book updates the current knowledge on the role of phytohormones in the control of plant growth and development, explores the mechanism responsible for the perception and signal transduction of phytohormones, and also provides a further understanding of the complexity of signal crosstalk and controlling downstream stress responses. There is next to none any book that provides update information on the phytohormones signi cance in tolerance to abiotic stress in plants. We extend our gratitude to all those who have contributed in making this book possible. Simultaneously, we would like to apologize unreservedly for any mistakes or failure to acknowledge fully. Aligarh, India Nafees A. Khan, Rahat Nazar, Noushina Iqbal Aveiro, Portugal Naser A. Anjum viPreface

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Contents 1 Signal Transduction of Phytohormones Under Abiotic Stresses....... 1 F. Eyidogan, M.T. Oz, M. Yucel, and H.A. Oktem 2 Cross Talk Between Phytohormone Signaling Pathways Under Both Optimal and Stressful Environmental Conditions............... 49 Marcia A. Harrison 3 Phytohormones in Salinity Tolerance: Ethylene and Gibberellins Cross Talk................................................................. 77 Noushina Iqbal, Asim Masood, and Nafees A. Khan 4 Function of Nitric Oxide Under Environmental Stress Conditions... 99 Marina Leterrier, Raquel Valderrama, Mounira Chaki, Morak Airaki, Jose "M. Palma, Juan B. Barroso, and Francisco J. Corpas 5 Auxin as Part of the Wounding Response in Plants................... 115 Claudia A. Casalongue ", Diego F. Fiol, Ramiro Par "s, Andrea V. Godoy, Sebastia "n D Ippo"lito, and Mar "a C. Terrile 6 How Do Lettuce Seedlings Adapt to Low pH Stress Conditions? A Mechanism for Low pH Induced Root Hair Formation in Lettuce Seedlings...................................................... 125 Hidenori Takahashi 7 Cytokinin Metabolism................................................... 157 Somya Dwivedi Burks 8 Origin of Brassinosteroids and Their Role in Oxidative Stress in Plants........................................................... 169 Andrzej Bajguz vii

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9 Hormonal Intermediates in the Protective Action of Exogenous Phytohormones in Wheat Plants Under Salinity..................... 185 Farida M. Shakirova, Azamat M. Avalbaev, Marina V. Bezrukova, Rimma A. Fatkhutdinova, Dilara R. Maslennikova, Ruslan A. Yuldashev, Chulpan R. Allagulova, and Oksana V. Lastochkina 10 The Role of Phytohormones in the Control of Plant Adaptation to Oxygen Depletion.................................................... 229 Vladislav V. Yemelyanov and Maria F. Shishova 11 Stress Hormone Levels Associated with Drought Tolerance vs. Sensitivity in Sun ower (Helianthus annuusL.)..................... 249 Cristian Ferna "ndez, Sergio Alemano, Ana Vigliocco, Andrea Andrade, and Guillermina Abdala 12 An Insight into the Role of Salicylic Acid and Jasmonic Acid in Salt Stress Tolerance.......................................... 277 M. Iqbal R. Khan, Shabina Syeed, Rahat Nazar, and Naser A. Anjum Index.......................................................................... 301 viiiContents

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Chapter 1 Signal Transduction of Phytohormones Under Abiotic Stresses F. Eyidogan, M.T. Oz, M. Yucel, and H.A. Oktem AbstractGrowth and productivity of higher plants are adversely affected by various environmental stresses which are of two main types, biotic and abiotic, depending on the source of stress. Broad range of abiotic stresses includes osmotic stress caused by drought, salinity, high or low temperatures, freezing, or ooding, as well as ionic, nutrient, or metal stresses, and others caused by mechanical factors, light, or radiation. Plants contrary to animals cannot escape from these environ mental constraints, and over the course of evolution, they have developed some physiological, biochemical, or molecular mechanisms to overcome effects of stress. Phytohormones such as auxin, cytokinin, abscisic acid, jasmonic acid, ethylene, salicylic acid, gibberellic acid, and few others, besides their functions during germination, growth, development, and owering, play key roles and coordinate various signal transduction pathways in plants during responses to environmental stresses. Complex networks of gene regulation by these phytohormones under abiotic stresses involve variouscis ortrans acting elements. Some of the transcrip tion factors regulated by phytohormones include ARF, AREB/ABF, DREB, MYC/ MYB, NAC, and others. Changes in gene expression, protein synthesis, modi ca tion, or degradation initiated by or coupled to these transcription factors and their correspondingcis acting elements are brie y summarized in this work. Moreover, crosstalk between signal transduction pathways involving phytohormones is explained in regard to transcriptional or translational regulation under abiotic stresses. F. Eyidogan (*) Baskent University, Ankara, Turkey e mail:[email protected] M.T. Oz M. Yucel H.A. Oktem Department of Biological Sciences, Middle East Technical University, Ankara, Turkey N.A. Khan et al. (eds.),Phytohormones and Abiotic Stress Tolerance in Plants, DOI 10.1007/978 3 642 25829 9_1, #Springer Verlag Berlin Heidelberg 20121

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1.1 Introduction Plants have successfully evolved to integrate diverse environmental cues into their developmental programs.Since theycannot escape from adverse constraints, they have been forced to counteract by eliciting various physiological, biochemi cal, and molecular responses. These responses include or lead to changes in gene expression, regulation of protein amount oractivity, alteration of cellular metab olite levels, and changes in homeostasis of ions. Gene regulation at the level of transcription is one of the major control points in biological processes, and transcription factors and regulators play key roles in this process. Phytohormones are a collection of trace amount growth regulators, comprising auxin, cytokinin, gibberellic acid (GA), abscisic acid (ABA), jasmonic acid (JA), ethylene, salicylic acid (SA), and few others (Tuteja and Sopory2008). Hormone responses are fundamental to the development and plastic growth of plants. Besides their regulatory functions during development, they play key roles and coordinate various signal transduction pathways during responses to environmental stresses (Wolters and J urgens2009). A range of stress signaling pathways have been elucidated through molecular genetic studies. Research on mutants, particularly ofArabidopsis, with defects in these and other processes have contributed substantially to the current understand ing of hormone perception and signal transduction. Plant hormones, such as ABA, JA, ethylene, and SA, mediate various abiotic and biotic stress responses. Although auxins, GAs, and cytokinins have been implicated primarily in developmental processes in plants, they regulate responses to stress or coordinate growth under stress conditions. The list of phytohormones is growing and now includes brassinosteroids (BR), nitric oxide (NO), polyamines, and the recently identi ed branching hormone strigolactone (Gray2004). Treatment of plants with exogenous hormones rapidly and transiently alters genome wide transcript pro les (Chapman and Estelle2009). InArabidopsis, hormone treatment for short periods (<1 h) alters expression of 10 300 genes, with roughly equal numbers of genes repressed and activated (Goda et al.2008; Nemhauser et al.2006; Paponov et al.2008). Not surprisingly, longer exposure to most hormones ( 1 h) alters expression of larger numbers of genes. Complex networks of gene regulation by phytohormones under abiotic stresses involve variouscis ortrans acting elements. Some of the transcription factors, regulators, and key components functioning in signaling pathways of phytohormones under abiotic stresses are described in this work. Moreover, changes in gene expression, protein synthesis, modi cation, or degradation initiated by or coupled to plant hormones are brie y summarized. 2 F. Eyidogan et al.

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