Shatz C J, Rakic P
J Comp Neurol. 1981 Feb 20;196(2):287-307. doi: 10.1002/cne.901960208.
The prenatal development of the cortical projections to the dorsal lateral geniculate nucleus (LGN), superior colliculus (SC) and pulvinar was studied by autoradiography of orthogradely transported 3H-proline injected into the occipital cortex of fetal rhesus monkeys aged from embryonic day 63 (E63) to E95. Differentiation of pyramidal neurons situated in the infragranular strata of the cortical plate (prospective layers 5 and 6, which give rise to these efferent projections) was also examined in Golgi preparations prepared from specimens of corresponding embryonic ages. In autoradiographs of the E63 fetus, no radioactive label was seen in subcortical structures. In two specimens injected around E70, label was present in the prospective magnocellular layers of the LGN and within the immediately surrounding cell-poor zones. At these young fetal ages, the presence of topographic order in the corticogeniculate projection could not be determined due to the large size of the injection sites relative to the small cerebral vesicles. By E84 the portion of the prospective parvocellular layers adjacent to the white matter also contained label which was characteristically wedge-shaped and appropriately located with respect to the site of the cortical injection, suggesting that topographic order is established. In the oldest fetus (E95) label in the LGN assumed a configuration similar to that seen in the adult. The cortical projection also invades the SC and pulvinar around E70. In the SC, label was initially confined to the stratum opticum, but by E84 it extended into the superficial gray. Thus, all known classes of efferent pathways from the visual cortex to subcortical structures are present by the middle of the 165-day gestational period in rhesus monkey. The one-month period, E63-E97, during which these efferent visual connections are established is characterized by the considerable growth and increased complexity of the dendritic arborization of pyramidal cells in the infragranular cortical layers of area 17. Thus the development of visual cortical efferents occurs in rough synchrony with the genesis of the afferent pathway from the LGN (Rakic, '76a; '79) and with the onset of morphological differentiation of pyramidal neurons in the infragranular cortical layers.
通过对妊娠63天(E63)至95天的恒河猴胎儿枕叶皮质注射正向运输的³H-脯氨酸进行放射自显影,研究了皮质向背外侧膝状核(LGN)、上丘(SC)和丘脑枕的投射在产前的发育情况。还对取自相应胚胎年龄标本的高尔基染色制剂中位于皮质板颗粒下层(即未来的第5层和第6层,这些传出投射由此产生)的锥体细胞的分化情况进行了检查。在E63胎儿的放射自显影片中,未在皮质下结构中看到放射性标记。在大约E70时注射的两个标本中,标记出现在LGN的未来大细胞层以及紧邻的细胞稀少区域内。在这些胎儿早期,由于注射部位相对于小的脑泡来说较大,所以无法确定皮质膝状体投射中是否存在拓扑顺序。到E84时,与白质相邻的未来小细胞层部分也含有标记,其形状呈典型的楔形,且相对于皮质注射部位位置恰当,这表明拓扑顺序已经建立。在最老的胎儿(E95)中,LGN中的标记呈现出与成年动物相似的形态。皮质投射在大约E70时也侵入了SC和丘脑枕。在SC中,标记最初局限于视层,但到E84时它扩展到了浅灰色层。因此,在恒河猴165天妊娠期的中期,所有已知的从视觉皮质到皮质下结构的传出通路都已存在。在E63 - E97这一个月期间,这些传出视觉连接得以建立,其特征是17区颗粒下层锥体细胞的树突分支有显著生长且复杂性增加。因此,视觉皮质传出纤维束的发育与来自LGN的传入通路的发生(拉基奇,1976a;1979)以及颗粒下层锥体细胞形态分化的开始大致同步。