Formation of the Spemann organizer in Xenopus laevis.

摘要

Spemann's Organizer, located in the dorsal marginal zone of the amphibian gastrula, induces and differentiates dorsal axial structures characteristic of this and other vertebrates. To trace the cellular origins of the Xenopus organizer, I labelled dorsal blastomeres of three of the four tiers (A, B, and C) of the 32-cell embryo with green, red, and blue fluorescent lineage tracers. A strong vegetal-ward displacement of labelled clones occurs between the late blastula and early gastrula stages but clones mix only slightly at their borders. The typical early gastrula organizer is composed of approximately 10% A1 progeny in its animal-most region, 70% B1 progeny in the central region, and 20% C1 progeny in vegetal and deep regions. Variability in the composition of the early gastrula organizer results from variability in positions of early cleavage planes and in pre-gastrulation movements. As the organizer involutes during gastrulation, forming dorsal axial mesoderm, clonal boundaries are greatly dispersed by cell intermixing. Within a clone, deep cells are displaced and intermixed more than superficial cells. The ventral marginal zone, composed mostly of C4 progeny, differs substantially from dorsal, and formation of ventral mesoderm occurs with less cell intermixing. Variability in blastomere progeny distributions within dorsal axial mesoderm of the late gastrula results mostly from variable intermixing of cells during gastrulation. Experiments to perturb later developmental events by molecular or embryonic manipulations at an early stage must take this variability into account along with the majority distributions of the fate map.;Within the early gastrula organizer, the genes Xbra, goosecoid, noggin and xNR3 are expressed differently in the animal-vegetal and superficial-deep dimensions. Using in situ hybridization combined with lineage labelling, I define distinct regions of the dorsal marginal zone. By the end of gastrulation, dorsal axial mesoderm cells derived from the organizer have altered their expression of the genes Xbra, goosecoid, noggin and xNR3 to suit their new positions in the anterior-posterior axis. At a given stage, a cell's position in the embryo rather than its lineage may be more important in determining which genes it will express.