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Not AvailablePlant RNA silencing systems are organized as a network, regulating plant developmental pathways and re-straining invading viruses, by sharing cellular components with overlapping functions. Host regulatory networks operate either at the transcriptional level via RNA-directed DNA methylation, or at the post-transcriptional stage interfering with mRNA to restrict viral infection. However, viral-derived proteins, including suppressors of RNAsilencing, favour virus establishment, and also affect plant developmental processes. In this investigation, wereport that Tomato leaf curl New Delhi virus-derived AC4 protein suppresses RNA silencing activity and mu-tational analysis of AC4 showed that Asn-50 in the SKNT-51 motif, in the C-terminal region, is a critical de-terminant of its RNA silencing suppressor activity. AC4 showed interaction with host AGO4 but not with AGO1,aggregated around the nucleus, and influenced cytosine methylation of the viral genome. The possible molecular mechanism by which AC4 interferes in the RNA silencing network, helps virus establishment, and affects plant development is discussed.Not Availabl