Axial, a zebrafish gene expressed along the developing body axis, shows altered expression in cyclops mutant embryos

Here, we report the cloning of a cDNA from zebrafish encoding a member of the fork head/HNF3 gene family. The gene, which we have called Axial, begins to be expressed just before gastrulation in a narrow region on the dorsal side of the embryo, the fish equivalent of the amphibian organizer. Expression can be detected in the involuted cells comprising the mesendoderm of the developing axis. At the end of gastrulation expression is turned on in the ventral neural plate in cells adjacent to the Axial-expressing mesodermal cells. Thus, Axial appears to be a target of both mesoderm induction and neural induction, leading to expression in cells of all three germ layers along the developing axis. Like the Brachyury gene. Axial is strongly induced by activin A, suggesting a role for endogenous activins in specifying the overlapping domains of expression of these two genes along the axis. Axial-expressing cells in the neuroectoderm include those of the future floor plate and cells of the ventral forebrain. In embryos homozygous for the cyclops mutation, expression is normal in mesendodermal cells but is absent from the ventral neural tube. The primary defects of cyclops mutants (lack of floor plate, deficiencies in the brain and cyclopia) correlate well with the expression domain of the Axial gene in wild-type neuroectoderm. The lack of Axial expression in cyclops neuroectoderm suggests that activation of Axial may be an immediate response of cyclops gene activity. Taken together, our data suggest that Axial plays a crucial role in specification of both the axial mesendoderm and the ventral central nervous system.