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Expected final online publication date for the Annual Review of Plant Biology, Volume 71 is April 29, 2020. Please see http//www.annualreviews.org/page/journal/pubdates for revised estimates.Cells confront DNA damage in every cell cycle. Among the most deleterious types of DNA damage are DNA double-strand breaks (DSBs), which can cause cell lethality if unrepaired or cancers if improperly repaired. In response to DNA DSBs, cells activate a complex DNA damage checkpoint (DDC) response that arrests the cell cycle, reprograms gene expres