Abstract: Actin is an ancient and abundant protein with well-established roles in fundamental processes ranging from cell migration to membrane transport. Most eukaryotic cells also contain at least eight actin-related proteins (ARPs) that are, themselves, conserved between organisms as divergent as yeast and mammals. Although many ARPs are cytoskeletal, recent biochemical and genetic work has demonstrated that some ARPs function largely or entirely in the nucleus. Evidence for the participation of both actin and ARPs in chromatin remodeling is becoming conclusive, and support for the still controversial involvement of actin in processes ranging from transcription to nuclear assembly is growing. The existence of conserved nuclear ARPs, together with accumulating biochemical, genetic and cell biology data, points to ancient and fundamental roles of actin in the nucleus, but the nature of these roles is just beginning to be revealed.