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Cellular dsRNAs are edited by adenosine deaminases that act on RNA (ADARs). While editing can alter mRNA-coding potential, most editing occurs in noncoding sequences, the function of which is poorly understood. Using dsRNA immunoprecipitation (dsRIP) and RNA sequencing (RNA-seq), we identified 1523 regions of clustered A-to-I editing, termed editing-enriched regions (EERs), in four stages of  Caenorhabditis elegans  development, often with highest expression in embryos. Analyses of small RNA-seq data revealed 22- to 23-nucleotide (nt) siRNAs, reminiscent of viral siRNAs, that mapped to EERs and were abundant in  adr-1;adr-2  mutant animals. Consistent with roles for these siRNAs in silencing, EER-associated genes (EAGs) were down-regulated in  adr-1;adr-2  embryos, and this was dependent on associated EERs and the RNAi factor RDE-4. We observed that ADARs genetically interact with the 26G endogenous siRNA (endo-siRNA) pathway, which likely competes for RNAi components; deletion of factors required for this pathway ( rrf-3  or  ergo-1 ) in  adr-1;adr-2  mutant strains caused a synthetic phenotype that was rescued by deleting antiviral RNAi factors. Poly(A) +  RNA-seq revealed EAG down-regulation and antiviral gene induction in  adr-1;adr-2;rrf-3  embryos, and these expression changes were dependent on  rde-1  and  rde-4 . Our data suggest that ADARs restrict antiviral silencing of cellular dsRNAs.

C. elegans ADARs antagonize silencing of cellular dsRNAs by the antiviral RNAi pathway