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      Some things get better with age: differences in salicylic acid accumulation and defense signaling in young and mature Arabidopsis

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          Abstract

          In Arabidopsis, much of what we know about the phytohormone salicylic acid (SA) and its role in plant defense comes from experiments using young plants. We are interested in understanding why young plants are susceptible to virulent strains of Pseudomonas syringae, while mature plants exhibit a robust defense response known as age-related resistance (ARR). SA-mediated signaling is important for defense in young plants, however, ARR occurs independently of the defense regulators NPR1 and WHY1. Furthermore, intercellular SA accumulation is an important component of ARR, and intercellular washing fluids from ARR-competent plants exhibit antibacterial activity, suggesting that SA acts as an antimicrobial agent in the intercellular space. Young plants accumulate both intracellular and intercellular SA during PAMP- and effector-triggered immunity, however, virulent P. syringae promotes susceptibility by suppressing SA accumulation using the phytotoxin coronatine. Here we outline the hypothesis that mature, ARR-competent Arabidopsis alleviates coronatine-mediated suppression of SA accumulation. We also explore the role of SA in other mature-plant processes such as flowering and senescence, and discuss their potential impact on ARR.

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          Isochorismate synthase is required to synthesize salicylic acid for plant defence.

          Salicylic acid (SA) mediates plant defences against pathogens, accumulating in both infected and distal leaves in response to pathogen attack. Pathogenesis-related gene expression and the synthesis of defensive compounds associated with both local and systemic acquired resistance (LAR and SAR) in plants require SA. In Arabidopsis, exogenous application of SA suffices to establish SAR, resulting in enhanced resistance to a variety of pathogens. However, despite its importance in plant defence against pathogens, SA biosynthesis is not well defined. Previous work has suggested that plants synthesize SA from phenylalanine; however, SA could still be produced when this pathway was inhibited, and the specific activity of radiolabelled SA in feeding experiments was often lower than expected. Some bacteria such as Pseudomonas aeruginosa synthesize SA using isochorismate synthase (ICS) and pyruvate lyase. Here we show, by cloning and characterizing an Arabidopsis defence-related gene (SID2) defined by mutation, that SA is synthesized from chorismate by means of ICS, and that SA made by this pathway is required for LAR and SAR responses.
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            Leaf senescence.

            Leaf senescence constitutes the final stage of leaf development and is critical for plants' fitness as nutrient relocation from leaves to reproducing seeds is achieved through this process. Leaf senescence involves a coordinated action at the cellular, tissue, organ, and organism levels under the control of a highly regulated genetic program. Major breakthroughs in the molecular understanding of leaf senescence were achieved through characterization of various senescence mutants and senescence-associated genes, which revealed the nature of regulatory factors and a highly complex molecular regulatory network underlying leaf senescence. The genetically identified regulatory factors include transcription regulators, receptors and signaling components for hormones and stress responses, and regulators of metabolism. Key issues still need to be elucidated, including cellular-level analysis of senescence-associated cell death, the mechanism of coordination among cellular-, organ-, and organism-level senescence, the integration mechanism of various senescence-affecting signals, and the nature and control of leaf age.
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              Salicylic Acid biosynthesis and metabolism.

              Salicylic acid (SA) has been shown to regulate various aspects of growth and development; it also serves as a critical signal for activating disease resistance in Arabidopsis thaliana and other plant species. This review surveys the mechanisms involved in the biosynthesis and metabolism of this critical plant hormone. While a complete biosynthetic route has yet to be established, stressed Arabidopsis appear to synthesize SA primarily via an isochorismate-utilizing pathway in the chloroplast. A distinct pathway utilizing phenylalanine as the substrate also may contribute to SA accumulation, although to a much lesser extent. Once synthesized, free SA levels can be regulated by a variety of chemical modifications. Many of these modifications inactivate SA; however, some confer novel properties that may aid in long distance SA transport or the activation of stress responses complementary to those induced by free SA. In addition, a number of factors that directly or indirectly regulate the expression of SA biosynthetic genes or that influence the rate of SA catabolism have been identified. An integrated model, encompassing current knowledge of SA metabolism in Arabidopsis, as well as the influence other plant hormones exert on SA metabolism, is presented.
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                Author and article information

                Contributors
                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                09 January 2015
                2014
                : 5
                : 775
                Affiliations
                [1]Department of Biology, McMaster University , Hamilton, ON, Canada
                Author notes

                Edited by: Hua Lu, University of Maryland at Baltimore County, USA

                Reviewed by: Saskia C. M. Van Wees, Utrecht University, Netherlands; Dale R. Walters, Scottish Agricultural College, Scotland

                *Correspondence: Robin K. Cameron, Department of Biology, McMaster University, 1280 Main Street West, Hamilton, ON L8S 4K1, Canada e-mail: rcamero@ 123456mcmaster.ca
                These authors have contributed equally to this work.

                This article was submitted to Plant-Microbe Interaction, a section of the journal Frontiers in Plant Science.

                Article
                10.3389/fpls.2014.00775
                4288333
                25620972
                28118def-d26a-4fe3-b066-ae9b1b4f91f6
                Copyright © 2015 Carella, Wilson and Cameron.

                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
                : 15 October 2014
                : 15 December 2014
                Page count
                Figures: 2, Tables: 0, Equations: 0, References: 55, Pages: 6, Words: 5347
                Categories
                Plant Science
                Perspective Article

                Plant science & Botany
                arabidopsis thaliana,pseudomonas syringae pv. tomato,age-related resistance,salicylic acid,antimicrobial,flowering,senescence,intercellular space

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