Postnatal development of corticospinal axon terminal morphology in the cat

The corticospinal system undergoes important postnatal development, leading to the mature topography and specificity of connections. The purpose of this study was to determine the time-course of development of corticospinal axonal branching and varicosity density within the cervical gray matter. Corticospinal neurons were labeled after small injections of the anterograde tracer biotinylated dextran amine into the primary motor cortex of cats. Tracer injection and transport times were adjusted to examine labeling at 25, 35, 55, and 75 days and in adults. We measured the numbers and lengths of nonreconstructed terminal and preterminal branches and the numbers and locations of axon varicosities. We found significant age-dependent increases in all morphologic measures. At 25 days, corticospinal axon branching was sparse, with only a few scattered varicosities. By day 35, the mean number of branches, varicosities per branch, and varicosity density increased. Several morphologic measures did not increase between day 35 and 55, but further changes occurred between 55 days and maturity. Beginning around day 55, there was extensive development of small terminal axon branches with high densities of varicosities. We also found, by using spatial point analysis, that there was an age-dependent increase in varicosity clustering. Our results show for the first time that terminal and preterminal corticospinal axon branches increase in complexity during a protracted early postnatal period. This developmental period extended beyond the early postnatal period of activity-dependent refinement of the topography of terminations. Comparison with the time-course of maturation of the cortical motor representation revealed development of substantial, albeit incomplete, branching and varicosity density of CS axons before cortical motor circuits effectively drive their spinal targets.