ADP-ribosylation factors (ARFs) are a family of Ras-related small GTPases, which are ubiquitously expressed and known to regulate many cellular processes including intracellular trafficking and membrane-related processes (D’Souza-Schorey and Chavrier 2006; Donaldson and Jackson 2011). Out of the six-member ARFs (ARF1–ARF6) in mammalian cells, ARF1 and ARF6 are well characterized. ARF1 regulates membrane trafficking between the endoplasmic reticulum (ER) and the Golgi, while ARF6 has been shown to modulate clathrin-dependent and independent endocytosis, exocytosis, and organization of actin cytoskeleton at the plasma membrane (Donaldson and Honda 2005). Like other Ras-related small GTPases, the ARF family of GTPases acts as binary switches by cycling between an inactive GDP-bound and an active GTP-bound conformations. The active/inactive cycle of ARFs is tightly...
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