The ontogenetic development of the serotoninergic system in the embryonic as well as in posthatching chick brain was studied with an indirect immunohistochemical technique with the aid of a specific antibody to serotonin (5-hydroxytryptamine). By embryonic day 4, rostral and caudal groups of serotonin-immunoreactive cell populations appeared in the mesencephalon and rostral and caudal rhombencephalon. At this stage, the rostral group had a considerable number of labelled cells that sent axons toward more rostral parts of brain, whereas the caudal group consisted of a small number of scattered serotonin-immunoreactive cells. The number of serotonin-positive cells increased with development, such that by embryonic day 8 almost all the serotoninergic cell groups found in the adult chick were already present. Serotoninergic-positive cells appeared in the paraventricular organ of the diencephalon as early as embryonic day 10. Judging from the cytoarchitectural organization of serotonin-immunoreactive cells, all of the serotoninergic cell groups in the chick brain seemed to be fully developed by embryonic day 16. On embryonic day 4, serotonin-immunoreactive fibers were found to enter into the marginal layer of the mesencephalon. Subsequently, serotonin-positive fibers ascended in the marginal layer of the brainstem up to the levels of the diencephalon and to the telencephalon on embryonic day 6 and 8, respectively. Serotonin-positive fibers, which first began to penetrate into the mantle layer on embryonic day 8, reached to the rostral pole of the telencephalon by embryonic day 10. In general, serotonin fibers were found in almost all brain regions by embryonic day 16. However, "terminal formation" in some nuclei did not seem to begin until the late embryonic or posthatching period. These observations indicate that the initial development of serotoninergic cell groups occurs during the first half of the 20th day of the incubation period of the chick. However, a longer time, ranging from early embryonic to posthatching stages, is necessary for the complete development of the serotoninergic projections.
Developmental changes of serotonin (5-hydroxytryptamine) neurons and fibers in the spinal cord of the embryo and posthatching chick were studied with immunohistochemical techniques with the aid of an antibody against serotonin. The first serotonin-immunoreactive fibers were found in the marginal layer of the cervical and lumbar spinal cord on embryonic days 6 and 8, respectively. There was a time lag of a few days between the first appearance of serotonin fibers in the marginal layer (embryonic days 6-8) and the time of penetration of serotonin fibers into the mantle layer (embryonic day 8 or older). The developments of serotonin innervation in the rostral parts of the spinal cord precedes that of caudal regions. Serotonin fibers penetrating into the mantle layer of the lumbar spinal cord were first found in lamina VII on embryonic day 8, whereas there were no serotonin-immunoreactive fibers in lamina IX by embryonic day 10. Large differences were found between embryonic day 16 and posthatching day 5 with regard to the density of serotonin varicosities and fibers in lamina IX, where profiles of soma and large-sized dendrites were heavily covered with varicosities. Laminae I and II first received serotonin fibers on embryonic day 16 and had a much denser innervation by posthatching day 5. There were no traces of serotonin fibers in lamina III in the stages examined up to posthatching day 5. Serotonin fibers were located in the lateral and ventral marginal layers in all specimens examined; only a few fibers were found in the dorsal marginal layer. Although few, serotonin-immunoreactive cell bodies were found in an area around the central canal of all animals from embryonic day 8 to adult. Some of these were located in the ependymal layer and sent processes toward the central canal; there were a small number of cells with long, fine processes. Serotonin-immunoreactive fibers in the spinal cord were not altered in regions rostral to the spinal transection, whereas all the serotoninergic fibers of the supraspinal origin were eliminated in the spinal cord caudal to the gap.
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