Microtubule transport and assembly cooperate to generate the microtubule array of growing axons

MTs are major architectural elements in growing axons. MTs overlap with each other along the axon, forming an array that is continuous from the cell body to the tip of the axon. The MT array constitutes a scaffolding that mechanically supports the elongate shape of the axon and also contributes directly to its shape. MTs also direct the transport of vesicular organelles between the cell body and the axon, and thereby determine, in part, the composition of the axon. In this article, I have discussed mechanisms involved in the elaboration of the MT array in growing axons, and I have emphasized the distinct but complementary roles of polymer transport mechanisms and local assembly dynamics. MTs for the axon originate in the cell body, and they are delivered to the axon by the polymer transport mechanisms. These mechanisms thus contribute directly to the shape of the axon by supplying it with essential architectural elements. The shape of the axon is further modulated by dynamic processes that alter cytoskeletal structure locally along its length. These dynamic processes include the assembly/disassembly mechanisms which influence polymer length and possibly number locally along the axon by subunit exchange between the monomer and polymer pools. In addition, the polymer transport mechanisms themselves are subject to modulation along the axon, as demonstrated by the observation that transport rate of MTs varies along the length of individual axons (Reinsch et al., 1991). Such local variations can, in and of themselves, change the number of MTs along the axon, and thereby focally affect axon shape. Thus, the dynamic processes of polymer transport and local assembly act cooperatively to shape the MT array of the axon, and thereby contribute directly to the elaboration of axonal morphology.