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      How phloem-feeding insects face the challenge of phloem-located defenses

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          Abstract

          Due to the high content of nutrient, sieve tubes are a primary target for pests, e.g., most phytophagous hemipteran. To protect the integrity of the sieve tubes as well as their content, plants possess diverse chemical and physical defense mechanisms. The latter mechanisms are important because they can potentially interfere with the food source accession of phloem-feeding insects. Physical defense mechanisms are based on callose as well as on proteins and often plug the sieve tube. Insects that feed from sieve tubes are potentially able to overwhelm these defense mechanisms using their saliva. Gel saliva forms a sheath in the apoplast around the stylet and is suggested to seal the stylet penetration site in the cell plasma membrane. In addition, watery saliva is secreted into penetrated cells including sieve elements; the presence of specific enzymes/effectors in this saliva is thought to interfere with plant defense responses. Here we detail several aspects of plant defense and discuss the interaction of plants and phloem-feeding insects. Recent agro-biotechnological phloem-located aphid control strategies are presented.

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          Most cited references158

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          Fungal effector proteins.

          It is accepted that most fungal avirulence genes encode virulence factors that are called effectors. Most fungal effectors are secreted, cysteine-rich proteins, and a role in virulence has been shown for a few of them, including Avr2 and Avr4 of Cladosporium fulvum, which inhibit plant cysteine proteases and protect chitin in fungal cell walls against plant chitinases, respectively. In resistant plants, effectors are directly or indirectly recognized by cognate resistance proteins that reside either inside the plant cell or on plasma membranes. Several secreted effectors function inside the host cell, but the uptake mechanism is not yet known. Variation observed among fungal effectors shows two types of selection that appear to relate to whether they interact directly or indirectly with their cognate resistance proteins. Direct interactions seem to favor point mutations in effector genes, leading to amino acid substitutions, whereas indirect interactions seem to favor jettison of effector genes.
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            Silverleaf whitefly induces salicylic acid defenses and suppresses effectual jasmonic acid defenses.

            The basal defenses important in curtailing the development of the phloem-feeding silverleaf whitefly (Bemisia tabaci type B; SLWF) on Arabidopsis (Arabidopsis thaliana) were investigated. Sentinel defense gene RNAs were monitored in SLWF-infested and control plants. Salicylic acid (SA)-responsive gene transcripts accumulated locally (PR1, BGL2, PR5, SID2, EDS5, PAD4) and systemically (PR1, BGL2, PR5) during SLWF nymph feeding. In contrast, jasmonic acid (JA)- and ethylene-dependent RNAs (PDF1.2, VSP1, HEL, THI2.1, FAD3, ERS1, ERF1) were repressed or not modulated in SLWF-infested leaves. To test for a role of SA and JA pathways in basal defense, SLWF development on mutant and transgenic lines that constitutively activate or impair defense pathways was determined. By monitoring the percentage of SLWF nymphs in each instar, we show that mutants that activate SA defenses (cim10) or impair JA defenses (coi1) accelerated SLWF nymphal development. Reciprocally, mutants that activate JA defenses (cev1) or impair SA defenses (npr1, NahG) slowed SLWF nymphal development. Furthermore, when npr1 plants, which do not activate downstream SA defenses, were treated with methyl jasmonate, a dramatic delay in nymph development was observed. Collectively, these results showed that SLWF-repressed, JA-regulated defenses were associated with basal defense to the SLWF.
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              Avoiding effective defenses: strategies employed by phloem-feeding insects.

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                Author and article information

                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                29 August 2013
                2013
                : 4
                : 336
                Affiliations
                [1] 1Institute of Phytopathology and Applied Zoology, Centre for BioSystems, Land Use and Nutrition, Justus-Liebig-University Giessen, Germany
                [2] 2Institute of General Botany, Justus-Liebig-University Giessen, Germany
                Author notes

                Edited by: Gary A. Thompson, Pennsylvania State University, USA

                Reviewed by: Sylvie Dinant, Institut National de la Recherche Agronomique, France; Jyoti Shah, University of North Texas, USA; Tom Wilkinson, University College Dublin, Ireland

                *Correspondence: Torsten Will, Institute of Phytopathology and Applied Zoology, Centre for BioSystems, Land Use and Nutrition, Justus-Liebig-University, Heinrich-Buff-Ring 26-32, D-35392 Giessen, Germany e-mail: torsten.will@ 123456agrar.uni-giessen.de

                Torsten Will and Alexandra C. U. Furch have contributed equally to the article.

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

                Article
                10.3389/fpls.2013.00336
                3756233
                24009620
                1ecfc831-359c-4b9f-9936-7fffc431077b
                Copyright © Will, Furch and Zimmermann.

                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
                : 25 March 2013
                : 09 August 2013
                Page count
                Figures: 1, Tables: 1, Equations: 0, References: 189, Pages: 12, Words: 0
                Categories
                Plant Science
                Review Article

                Plant science & Botany
                aphid-plant interactions,aphid saliva,p-proteins,plant defense,phloem,sieve element occlusion

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