Properties of the dorsalizing signal in gastrulae of Xenopus laevis.

According to the 'three signal model', the regional specification of tissue type within the mesoderm of Xenopus laevis occurs in a process called 'dorsalization'. We have studied the timing and transmission characteristics of this signal, and assessed the dorsalizing activity of the lithium ion and a panel of cytokines.The marginal zone has been fate mapped during gastrulation by colloidal gold labelling and it is shown that the ventral tissue undergoes substantial circumferential expansion. The fate map information is used to provide tissues of constant cellular composition for experiments conducted at different stages.The stage at which dorsalization can occur has been investigated by means of heterochronic dorsal-ventral combinations. The results indicate that the interaction occurs during gastrulation, with a decline in both signal strength and competence of the ventral marginal zone to respond as gastrulation proceeds.The signal is capable of passing through arrangements of membranes that exclude the possibility of cytoplasmic contact, implying that it can be carried by a diffusible morphogen.The effect on the ventral marginal zone of lithium and a number of cytokines has also been studied. While none appears to function as a dorsalizing signal, lithium acts during blastula stages to alter the response to the mesoderm-inducing signal such that the inductions are of a more dorsal character.These data confirm that the dorsalizing signal is independent of and operates later than the signal(s) from the vegetal hemisphere that induce mesoderm during the blastula stages.