The septins are a family of GTPase enzymes, some of which are required for cytokinesis and others of which are associated with exocytosis. Members of the septin family can form heteropolymer complexes and also play a role in the organization of new growth in organisms. The transcriptional regulation of all septins is complex, resulting in alternatively spliced variants. At least three septins (Septin 1, 2 and 4) are associated with a Tau-based paired helical filament core and may contribute to the formation of neurofibrillary tangle as integral constituents of paired helical filaments. Septin 3 (G-Septin), a GTP-binding protein, is highly expressed in brain and is regulated by protein kinase G in neurons. The human SEPT4 (H5/PNUTL2/CDCrREL-2) gene encodes ARTS (for apoptosis-related protein in the TGFbeta signaling pathway), which is expressed in many cells and acts to enhance cell death induced by TGFbeta or, to a lesser extent, by other apoptotic agents. ARTS is localized to mitochondria and translocates to the nucleus when apoptosis occurs.