The development of the septal region in the rat. II. Morphogenesis in normal and X-irradiated embryos

Abstract

Morphogenesis of the septal region was examined in normal rat embryos from embryonic day (E) 10 to E22. The greater part of the septal region is postulated to form from two separate anlagen which can be clearly distinguished in the telencephalon by E13 and E14. One lies in the anterior ventromedial wall and presumably forms the nucleus of the diagonal band, medial, lateral, and triangular septal nuclei. The other lies in the posterior ventrolateral ridge and presumably forms the bed nuclei of the stria terminalis and the anterior commissure. On E15, the early differentiating cells in these anlagen fuse in the same region where the anterior commissure will cross on E17. With later embryonic development, differentiating cells of the strial bed nucleus accumulate rostral and caudal to the fused area. The same pattern is found in the medial and triangular septal nuclei and in the nucleus of the diagonal band. The differentiating cells of the lateral septal nucleus accumulate dorsal and lateral to the medial and triangular septal nuclei. On E16 and E17, a prominent subependymal zone develops in the anterior septal region and presumably gives rise to the nucleus accumbens. A quantitative analysis was made of three cell zones (neuroepithelium, subependymal zone, differentiating cell zone) at coronal levels through the developing nucleus accumbens and the nucleus of the diagonal band (anterior level) and the medial and lateral septal nuclei (middle and posterior levels). At all levels, the area of the neuroepithelium continually declines, that of the differentiating cell zone continually increases, and that of the subependymal zone shows a rise and decline. On a proportional basis, both the neuroepithelium and subependymal zone occupy significantly more area anteriorly than posteriorly, while the differentiating cell zone shows the reverse gradient. To accurately locate regions of primitive mitotic and migratory cells within the zones at each level, the number of cells surviving a single exposure to 200 R X-rays in embryonic brains (E15-E22) were compared with controls. Each zone responded differently to X-ray insult. The radiosensitivity of the neuroepithelium decreases significantly after E19; the subependymal zone is highly radiosensitive throughout; the differentiating cell zone is radioresistant throughout. The significance of these findings is discussed in the light of the autoradiographic determination of the time of formation of septal neurons (Bayer, '79).