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Some aspects of this disclosure provide compositions, methods, and kits for improving the specificity of RNA-programmable endonucleases, such as Cas9. Also provided are variants of Cas9, e.

Also provided are compositions, methods, and kits for site-specific nucleic acid modification using Cas9 fusion proteins e. Such Cas9 variants are useful in clinical and research settings involving site- specific modification of DNA, for example, genomic modifications.

In a variety of organisms, including mammals, site-specific endonucleases have been used for genome engineering by stimulating either non-homologous end joining or homologous recombination. In addition to providing powerful research tools, site-specific nucleases also have potential as gene therapy agents, and two site-specific endonucleases have recently entered clinical trials: l CCR, targeting a human CCR-5 allele as part of an anti-HIV therapeutic approach NCT, NCT,.

For example, imperfect specificity of engineered site-specific binding domains has been linked to cellular toxicity and undesired alterations of genomic loci other than the intended target. Most nucleases available today, however, exhibit significant off-target activity, and thus may not be suitable for clinical applications. An emerging nuclease platform for use in clinical and research settings are the RNA-guided nucleases, such as Cas9. Technology for engineering nucleases with improved specificity is therefore needed.