Restoration of the organizer after radical ablation of Hensen's node and the anterior primitive streak in the chick embryo.

Abstract

The region of the amniote embryo corresponding to Spemann's organizer in amphibians is Hensen's node, which lies at the tip of the primitive streak during gastrulation. It is a special site in the embryo that can be defined by the presence of progenitors of several axial tissues (notochord, prechordal mesoderm, somites, gut endoderm), by characteristic cell movements, by specific patterns of gene expression (e.g. goosecoid, HNF-3beta, Sonic hedgehog) and, most importantly, by its ability to induce a complete axis, including host-derived neural tissue, when transplanted to an ectopic site. Here, we show that complete removal not only of the node but also of the anterior 40% of the primitive streak leads to the development of normal embryos containing cells with all the fates normally produced by the node. Cell movement pathways through the regenerated node are identical to those seen in the normal embryo. The patterns of expression of HNF-3beta and Sonic hedgehog are also restored, as is their left/right asymmetry, but goosecoid expression is not. When the regenerated node is transplanted to an ectopic site, it induces a complete embryonic axis that includes a fully patterned, host-derived central nervous system. Analysis of the properties of cells surrounding the site of ablation shows that they acquire these properties gradually. We suggest that the organizer is a region of the embryo that is defined by cell interactions and that the node normally inhibits the organizer state in neighbouring cells.