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-catenin/XTcf-3 complex binds to the
siamois promoter to regulate dorsal axis specification
in Xenopus
1 Department of Biochemistry and
2 Howard Hughes
Medical Institute and Department of Pharmacology, University of
Washington School of Medicine, Seattle, Washington 98195-7350 USA
The Wnt pathway regulates the early dorsal-ventral axis
in Xenopus through a complex of 
-catenin and HMG box
transcription factors of the Lef/Tcf family. We show that
the promoter of the dorsalizing homeo box gene siamois is a
direct target for the -catenin/XTcf-3 complex,
establishing a link between the Wnt pathway and the activation
of genes involved in specifying the dorsal axis. By injecting
siamois reporter constructs into the animal pole of
Xenopus embryos, we show that a 0.8-kb fragment of the
siamois promoter is strongly activated by -catenin. The proximal 0.5 kb, which is also activated by -catenin, contains three
Lef/Tcf-binding sites. Mutations in these sites eliminate the -catenin-mediated activation of siamois and show that
siamois is regulated by the -catenin/XTcf-3
complex, in combination with additional transcriptional activators.
When expressed at the equator of the embryo, the siamois
promoter is activated to much higher levels on the dorsal side than the
ventral side. Ectopic ventral expression of -catenin raises the
ventral expression of the siamois promoter to the dorsal
levels. Conversely, ectopic dorsal expression of dominant-negative
XTcf-3 abolishes the dorsal activation of the siamois promoter.
Furthermore, elimination of the Lef/Tcf sites elevates
the ventral expression of siamois, revealing a repressive role
for XTcf-3 in the absence of -catenin. Finally, we find that the
endogenous siamois activator, although present throughout the
dorsal side of the embryo, is most potent in the dorsal vegetal region.
We propose that the dorsal activation of siamois by the
-catenin/XTcf-3 complex combined with the ventral repression of siamois by XTcf-3 results in the restriction of endogenous siamois expression to the dorsal side of
Xenopus embryos.
[Key Words: Xenopus; homeobox; HMG box; dorsoventral patterning; Wnt; Lef/Tcf; siamois]
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