The Cas9 protein, derived from the Type II CRISPR (clustered, regularly interspaced short palindromic repeat) bacterial immune systems, is emerging as a powerful tool for engineering the genome in many organisms. Since its discovery, Cas9-based technology has been adopted for editing, regulating, and imaging the genome, as well as rewriting the epigenetic status, all in a sequence-specific manner. With all of these advances, we have just begun to explore its possible applications. Here, we describe current models for Cas9 function based on structural and biochemical studies. We focus on its applications in genome editing, regulation, and imaging, discuss other possible applications and some technical considerations, and highlight the many advantages that CRISPR-Cas9 technology offers. Expected final online publication date for the Annual Review of Biochemistry Volume 85 is June 2, 2016. Please see http://www.annualreviews.org/catalog/pubdates.aspx for revised estimates.
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