Müller F, O'Rahilly R
School of Medicine, University of California, Davis, Calif, USA.
Cells Tissues Organs. 2004;177(1):2-20. doi: 10.1159/000078423.
The caudal region of the trunk was reassessed in 52 serially sectioned human embryos of stages 8-23, 42 of which were controlled by precise graphic reconstructions. The following observations, new for the human, are presented. (1) The neurenteric canal is an important landmark because rostral to it the neural plate of stages 8, 9, and the main part of the notochord develop, whereas caudal to it the neural plate of stages 10-12 and the caudal portion of the notochord are formed. All somites at stages 9-11 and probably also at stage 12 arise rostral to the site of the neurenteric canal. (2) A 'chordoneural hinge' can be detected in stages 10 and 11, where the caudal part of the neural plate gives off cells that probably participate in the production of mesenchyme. (3) When apparent disappearance of the epiblast is used as a criterion, then the primitive streak seems to end during stage 9. (4) The caudal eminence, derived from the primitive streak and covered by ectoderm, forms at stage 10 caudal to the site of the former neurenteric canal and persists as a terminal cap to at least stage 14, although formation of mesenchyme continues in stages 15 to 17 or 18. (5) As the region rostral to the site of the neurenteric canal grows because of the development of somites, the caudal eminence is shifted caudally. (6) The caudal eminence is most active developmentally during stage 13, when most of the required (ca 6 out of 9) pairs of somites appear. (7) The eminence produces the caudal part of the notochord and, after closure of the caudal neuropore, all caudal structures, but it does not produce even a temporary 'tail' in the human. (8) A temporal overlap results between primary and secondary development in the caudal part of the notochord. (9) Primary development begins very early with the formation of the inner cell mass at stage 3, and includes the development of the somites rostral to the neurenteric canal, whereas secondary development, with the exception of the notochord caudally, commences at stage 12. (10) Primary neurulation lasts from stage 8 to stage 12, secondary from stage 12 to stages 17 or 18. (11) Secondary development and secondary neurulation are characterized morphologically by direct formation of structures (notochord, postcloacal gut, neural cord/neural tube) from mesenchyme.
对52个8 - 23期连续切片的人类胚胎的躯干尾部区域进行了重新评估,其中42个通过精确的图形重建进行对照。以下是关于人类的新观察结果。(1)神经肠管是一个重要的标志,因为在其头侧,8、9期的神经板和脊索的主要部分发育形成,而在其尾侧,10 - 12期的神经板和脊索的尾端部分形成。9 - 11期以及可能12期的所有体节都在神经肠管部位的头侧产生。(2)在10和11期可检测到一个“脊神经铰链”,神经板的尾端部分在此处释放细胞,这些细胞可能参与间充质的产生。(3)当以上胚层明显消失作为标准时,原条似乎在9期结束。(4)尾隆凸源自原条,被外胚层覆盖,在10期于原神经肠管部位的尾侧形成,并至少持续到14期作为终帽存在,尽管间充质的形成在15至17或18期仍在继续。(5)由于体节的发育,神经肠管部位头侧的区域生长,尾隆凸向尾侧移位。(6)尾隆凸在13期发育最为活跃,此时大部分所需的(9对中的约6对)体节出现。(7)尾隆凸产生脊索的尾端部分,在尾神经孔闭合后,产生所有尾侧结构,但在人类中它甚至不会产生一个临时的“尾巴”。(8)脊索尾端部分的初级和次级发育之间存在时间上的重叠。(9)初级发育在3期内细胞团形成时就很早就开始了,包括神经肠管头侧体节的发育,而次级发育,除了尾侧的脊索外,在12期开始。(10)初级神经胚形成从8期持续到12期,次级神经胚形成从12期持续到17或18期。(11)次级发育和次级神经胚形成在形态学上的特征是由间充质直接形成结构(脊索、泄殖腔后肠、神经索/神经管)。