Cell Defense Responses Associated with Localized and Systemic Resistance to Phytophthora parasitica Induced in Tomato by an Arbuscular Mycorrhizal Fungus

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Abstract
The arbuscular mycorrhizal fungus Glomus mosseae is able to confer bioprotection against Phytophthora parasitica in tomato roots. Localized and induced systemic resistance (ISR) have been demonstrated to be involved in pathogen control in mycorrhizal and nonmycorrhizal roots with a split root experimental system. Decreased pathogen development in mycorrhizal and nonmycorrhizal parts of mycorrhizal root systems is associated with accumulation of phenolics and plant cell defense responses. G. mosseae-containing cortical cells in the mycorrhizal tissues are immune to the pathogen and exhibit a localized resistance response with the formation of cell wall appositions reinforced by callose adjacent to intercellular hyphae. The systemically induced resistance in nonmycorrhizal root parts is characterized by elicitation of host wall thickenings containing non-esterified pectins and PR-1a protein in reaction to intercellular pathogen hyphae, and by the formation of callose-rich encasement material around P. parasitica hyphae that are penetrating root cells. PR-la protein is detected in the pathogen wall only in these tissues. None of these cell reactions are observed in nonmycorrhizal pathogen-infected root systems, where disease development leads to host cell death. The cellular and molecular basis of bioprotection by an arbuscular mycorrhizal fungus is discussed in relation to that induced by other nonpathogenic microorganisms.