Schoenwolf G C, Delongo J
Am J Anat. 1980 May;158(1):43-63. doi: 10.1002/aja.1001580106.
Formation of the future lumbosacral level of the spinal cord was studied in two-day-old chick embryos by light and electron (transmission and scanning) microscopy. A neurulation overlap zone occupied this level. The dorsal portion of the neural tube formed by bending of the neural plate and approximation and fusion of neural folds (i.e., by primary neurulation), and the ventral part formed during secondary neurulation by cavitation of an initially solid, compact mass of cells, the medullary cord, derived from the tail bud. Secondary neurulation involved four morphogenetic processes: (1) segregation of the cells of the prospective medullary cord from cells of adjacent regions, (2) formation of a precisely delimited medullary cord, (3) cavitation of the central portion of this cord, and (4) coalescence of all lumina into a single, central cavity. Cell segregation was associated with the formation of a layer of primarily extracellular materials between adjacent organ rudiments. The source and composition of these materials are unknown. Formation of the medullary cord entailed considerable elongation of the peripheral cells of this developing structure and the fabrication of small intercellular juctions, first at the basal (outer) ends of the elongating peripheral cells, and then at their apical (inner) ends. These arranged, columnar cells, having characteristics similar to those of the neural plate, and an inner cluster of irregularly shaped and arranged cells. Cavitation always occurred first at the junction between these two cellular populations. The central cells of the medullary cord also eventually elongated, like the peripheral cells, and may have been intercalated into the lateral walls of the developing neural tube as lumina coalesced.
通过光学显微镜以及电子显微镜(透射电镜和扫描电镜)对2日龄鸡胚未来脊髓腰骶段的形成进行了研究。该水平区域存在神经胚形成重叠区。神经管的背侧部分由神经板弯曲以及神经褶的靠近和融合形成(即通过原发性神经胚形成),而腹侧部分在继发性神经胚形成过程中,由最初坚实紧密的细胞团块——髓索(源自尾芽)空化形成。继发性神经胚形成涉及四个形态发生过程:(1)预期髓索的细胞与相邻区域的细胞分离;(2)形成精确界定的髓索;(3)该髓索中央部分空化;(4)所有管腔合并成一个单一的中央腔。细胞分离与相邻器官原基之间主要由细胞外物质构成的一层结构的形成有关。这些物质的来源和组成尚不清楚。髓索的形成需要该发育结构的外周细胞大幅伸长,并形成小的细胞间连接,首先在伸长的外周细胞的基部(外侧)末端形成,然后在其顶端(内侧)末端形成。这些排列成柱状的细胞具有与神经板相似的特征,以及一群内部形状和排列不规则的细胞。空化总是首先发生在这两个细胞群体之间的交界处。髓索的中央细胞最终也像外周细胞一样伸长,并且在管腔合并时可能插入到发育中的神经管的侧壁中。