Intracellular transport in axonal microtubular domains II. Velocity profile and energetics of circumtubular flow

Abstract

The microtubule is a highly efficient vectorial structure that could orient a transport force generating mechanism and also absorb the recoil produced by vectorial force generation. We have assumed that a nonspecific shear force is generated in a narrow annulus around the microtubule and have calculated the velocity profiles in the shear flow and drag flow regions that result from such a mechanism. This circumtubular flow of low visocosity cytoplasm is thought to be the basic carrier stream that produces the observed axoplasmic transport phenomena. These carrier streams are devoid of neurofilaments and form the halos or exclusion zones seen around microtubules in electron micrographs. Individual carrier streams may merge hydrodynamically to produce transport domains that are capable of moving large organelles in a saltatory manner. Exchange of material between the low viscosity transport domains and the high macroviscosity neurofilament regions produces mass fluxes akin to those found in chromatographic columns. Calculations of energy required to maintain streaming and of the energy available to the transport system show a close correspondence and demonstrate that a continuous carrier stream activity is energetically feasible.