A quantitative analysis of axon outgrowth, axon loss, and myelination in the rat pyramidal tract

Abstract

A quantitative analysis of the development of the pyramidal tract (PT) was carried out at the level of the caudal medulla oblongata and at the sixth cervical spinal segment (C 6), in rats ranging in age from embryonic day 20 (E20) to the adult of 90 days postnatally (P90). The axon number in the right medullary PT rises from 27,000 axons at E20 to 391,000 axons at P4. Growth cones are abundant during this period, but can still be observed occasionally at P7. After P4, the axon number is reduced by 62%, to 150,000 in the adult. A rapid axon loss until P14 is followed by a gradual axon loss, continuing beyond the third postnatal week. A similar biphasic axon loss was observed in the cervical PT. At P2 and at P7, concentrations of electron-dense material were observed in 0.5–0.7% of the axon profiles in the medullary PT. Since at P21 this feature was only observed in 0.2% of the axons, it might represent an early sign of axon loss. Myelination starts in the medullary PT at P7. Especially during the third postnatal week, the number of myelinated axons increases rapidly. In the adult rat PT, both at medullary and cervical levels, about one third of the axons are still unmyelinated. The results indicate that the development of the rat PT is characterized by a gradual outgrowth of its fibers and by a protracted, biphasic axon loss. Furthermore, comparing the PT at the medulla, at C 3 (ref. 18), and at C 6, a rostrocaudal decrease in axon number was observed during development as well as at the adult stage. Therefore, no evidence was found for increased axon branching in the tract in the cervical intumescence.