Conditioning of cellular defence responses to powdery mildew in cereal leaves by prior attack

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Abstract
Field‐grown plants sequentially encounter many different fungal pathogens and nonpathogens that are capable of triggering an array of responses that may affect the subsequent level of disease they develop following later pathogen attack. These changes, which are induced by prior encounters, may be manifest as increased susceptibility or enhanced resistance to later pathogen attack; they may be expressed systemically or their effects may be localized within a few cells distance of the original encounter site. Here, we review our recent investigations of cellular changes effected by sequential inoculations of cereal leaves with the powdery mildew fungus Blumeria graminis DC.In susceptible barley and oats, a successful B. graminis attack followed by haustorium formation, renders the attacked cell, and to some extent its adjacent cells, highly accessible to later B. graminis attacks. By contrast, a failed attack due to papilla formation by the attacked host cells, renders the attacked cell and its adjacent cells highly inaccessible to later B. graminis attacks. Importantly, barley carrying the mlo5 allele for powdery mildew resistance is also conditioned to accessibility if prior attacks by an mlo‐virulent isolate penetrates successfully. In the partial resistant oat cultivar Maldwyn B. graminis attacks either succeeded, failed due to papilla deposition, or failed because the attacked host cells died in response to the attacks. Sequential inoculation of Maldwyn demonstrated the induction of accessibility and inaccessibility, as well as a complete suppression of cell death response to attack where the cells had survived an earlier attack. Furthermore, when a prior attack induced cell death, a later attack on adjacent cells caused greatly increased rate of cell death, demonstrating potentiation of cell death. The importance of the induced cellular changes for plant resistance in the field is discussed.

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