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Cullin Ring E3 ligases(CRLs)are a large family of ubiquitin ligases mediating the degradation of~20%of the proteome.CRLs are activated by neddylation and regulated by the deneddylase COP9 Signalosome(CSN).CSN biochemically inhibits but genetically activates CRLs,underscoring the importance of dynamic complex assembly,which remains poorly understood.