Multiple roles for actin in secretory and endocytic pathways.

Actin filaments play multiple roles in the secretory pathway and in endosome dynamics in mammals, including maintenance of Golgi structure, release of membrane cargo from the trans-Golgi network (TGN), endocytosis, and endosomal sorting dynamics. In addition, TGN carrier transport and endocytosis both occur by multiple mechanisms in mammals. Actin likely plays a role in at least four mammalian endocytic pathways, five pathways for membrane release from the TGN, and three processes involving endosomes. Also, the mammalian Golgi structure is highly dynamic, and actin is likely important for these dynamics. One challenge for many of these processes is the need to deal with other membrane-associated structures, such as the cortical actin network at the plasma membrane or the matrix that surrounds the Golgi. Arp2/3 complex is a major actin assembly factor in most of the processes mentioned, but roles for formins and tandem WH2-motif-containing assembly factors are being elucidated and are anticipated to grow with further study. The specific role for actin has not been defined for most of these processes, but is likely to involve the generation of force for membrane dynamics, either by actin polymerization itself or by myosin motor activity. Defining these processes mechanistically is necessary for understanding membrane dynamics in general, as well as pathways that utilize these processes, such as autophagy.