Dorsalization and neural induction: properties of the organizer in Xenopus laevis.

We have studied the action of the organizer in Xenopus laevis using grafts labelled with horseradish peroxidase (HRP). Orthotropic grafts of the dorsal marginal zone (the organizer) from an HRP-labelled embryo into an unlabelled host showed that this region contributes to the anterior archenteron wall, to the entire craniocaudal extent of the notochord and to a few cells in the somites. Little or no contribution was made to the neural tube. Orthotopic grafts of the ventral marginal zone (the tissue that responds to a grafted organizer) indicated that it only contributes to the posterior half of the embryo. Within this region it spreads around the entire ventrolateral mesoderm, occasionally contributing a few cells to the somites. The posterior endoderm was also heavily labelled. When the dorsal marginal zone from an HRP-labelled embryo was inserted into a slit cut in the ventral marginal zone of an unlabelled host a mirror-symmetrical double-dorsal duplicated embryo resulted, in which only the notochord and a few cells in the somites of the secondary embryo were derived from the graft. The bulk of the secondary somites was, therefore, derived from host ventral marginal zone tissue which normally makes very little contribution to the somites. This indicates that host ventral marginal zone becomes dorsalized by the graft. The neural tube of the secondary embryo was also unlabelled, showing that it was induced by the influence of the graft on the overlying ectoderm, which normally forms ventral epidermis. We have also grafted ventral marginal zone tissue into a slit cut into the dorsal marginal zone of a host embryo. HRP-labelled tissue was grafted into an unlabelled embryo and vice versa. This graft did not produce a double ventral embryo and this reinforces the traditional view that the dorsal marginal zone is a special signalling region. Instead, the resulting embryos usually had a twinned notochord with the graft tissue in between, differentiated as somite. This confirms that juxtaposing ventral and dorsal marginal zone 'dorsalizes' the ventral tissue but does not affect the dorsal tissue which differentiates, as usual, as notochord. Thus, our results allow us to conclude that the organizer mediates two distinct interactions in bringing about the formation of duplicated embryos. The first is dorsalization of adjacent ventral mesoderm and the second is the induction of neuroepithelium from ectoderm overlying the new archenteron roof.