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4.2.3 USP1 A genome-wide CRISPR/Cas9 knockdown screen identified ubiquitin-specific protease 1 ( USP1 ) as a novel regulator that positively modulates BMAL1 protein levels ( USP1 , but not its catalytically inactive mutant, led to increased BMAL1 protein abundance, whereas USP1 knockdown via shRNA or CRISPR knockout significantly reduced BMAL1 levels in U2OS cells ( in vitro and in mouse tissues, further confirming its role in regulating BMAL1 stability ( BMAL1 mRNA levels