What we talk about when we talk about nuclear actin

Abstract

In the cytoplasm, actin filaments form crosslinked networks that enable eukaryotic cells to transport cargo, change shape, and move. Actin is also present in the nucleus but, in this compartment, its functions are more cryptic and controversial. If we distill the substantial literature on nuclear actin down to its essentials, we find four, recurring, and more-or-less independent, claims: (1) crosslinked networks of conventional actin filaments span the nucleus and help maintain its structure and organize its contents; (2) assembly or contraction of filaments regulates specific nuclear events; (3) actin monomers moonlight as subunits of chromatin remodeling complexes, independent of their ability to form filaments; and (4) modified actin monomers or oligomers, structurally distinct from canonical, cytoskeletal filaments, mediate nuclear events by unknown mechanisms. We discuss the evidence underlying these claims and as well as their strengths and weaknesses. Next, we describe our recent work, in which we built probes specific for nuclear actin and used them to describe the form and distribution of actin in somatic cell nuclei. Finally, we discuss how different forms of nuclear actin may play different roles in different cell types and physiological contexts.

Keywords: cytoskeleton; exportin 6; importin 9; nuclear actin; nuclear myosins; oocyte.

Figure 1. Proposed forms of actin in the nucleus. (A) Crosslinked networks of conventional actin filaments, (B) linear filaments that support motor-based transport, (C) monomers that regulate chromatin-based protein complexes and (D) non-canonical, modified actin monomers or oligomers.

Figure 2. A comparison of known nuclear actin architectures. Oocyte germinal vesicles contain cross-linked actin networks (A) that undergo myosin-based contraction following nuclear envelope breakdown (B). In somatic nuclei, short actin filaments and a functional monomeric pool are present (C) and reorganize into cofilin-actin rods (D) following environmental stress.