Apoplastic and vascular defences

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Abstract
The apoplast comprises the intercellular space between cell membranes, includes the xylem, and extends to the rhizoplane and the outer surfaces of the plant. The apoplast plays roles in different biological processes including plant immunity. This highly specialised space is often the first place where pathogen recognition occurs, and this then triggers the immune response. The immune response in the apoplast involves different mechanisms that restrict pathogen infection. Among these responses, secretion of different molecules like proteases, proteins related to immunity, small RNAs and secondary metabolites play important and often additive or synergistic roles. In addition, production of reactive oxygen species occurs to cause direct deleterious effects on the pathogen as well as reinforce the plant’s immune response by triggering modifications to cell wall composition and providing additional defence signalling capabilities. The pool of available sugar in the apoplast also plays a role in immunity. These sugars can be manipulated by both interactors, pathogens gaining access to nutrients whilst the plant's responses restrict the pathogen’s access to nutrients. In this review, we describe the latest findings in the field to highlight the importance of the apoplast in plant–pathogen interactions and plant immunity. We also indicate where new discoveries are needed.

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