Synthetic Priming of Transcriptional Memory

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Plants have evolved the capacity to form 'memories' of pathogen attacks and other environmental stresses. These memories enable plants to mount a more vigorous response when the stress is encountered again. This effect, known as 'priming', has been observed across eukaryotes. Consistent with the ability of chromatin to carry information across mitotic and sometimes even meiotic divisions, recent research has linked chromatin changes to priming. However, it is not known how such chromatin changes produce a more effective response. Given that rapid deployment of transcriptional programs is critical for overcoming stresses, an attractive hypothesis is that priming re-sculpts chromatin to adopt a transcriptionally poised state that facilitates re-activation of stress-response genes. The proposed work will test this hypothesis for pathogen priming in Arabidopsis thaliana and will then seek to engineer plants to be primed for pathogen attack. In preliminary work, I have identified several pathogen primed genes, and I have established new tools that will allow me to address previously inaccessible questions regarding the mechanism by which chromatin confers transcriptional memory. In Aim 1, I will reveal the chromatin signature of priming (genomic techniques). In Aim 2, I will identify the factors acting downstream to perceive primed chromatin (quantitative proteomics). In Aim 3, I will discover the machinery required for transcriptional memory (forward and reverse genetics). In Aim 4, I will use the insights gained from Aims 1-3 to ask which chromatin features are necessary and sufficient to establish pathogen-primed transcriptional memory (epigenome engineering). This research, at the interface of epigenetics and immunity, will provide fundamental knowledge of the chromatin mechanisms that perpetuate active gene states, and explores the potential to instil primed 'memories' into naïve plants.