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Fig. 6. Reduced Foxf gene dosage has pleiotropic effects on paracrine signaling in the intestinal mesenchyme: more Wnt and less Bmp. (A) Whole-mount in situ hybridization of E14.5 embryos with a Wnt5a probe. Expression is significantly higher in Foxf2^-/- mutant (right) than in wild-type (left) small intestine. (B-E) X-gal staining of E15.5 Bmp4^lacZ/+ embryos. Bmp4-driven ß-galactosidase activity is significantly reduced in the gastrointestinal tract of Foxf2^-/- (C), compared with wild type (B). The same embryos have comparable levels of staining in organs where Foxf2 is not expressed, such as eyelids and hair follicles (D,E). (F-J) Bmp signaling inhibits Wnt5a expression in gastrointestinal mesenchyme. Wnt5a whole-mount in situ hybridization of gastric (G,I) and intestinal (F,H,J) explants from Foxf2^-/- (F-I) or wild-type (J) E12.5 embryos cultured for 24 hours with beads (arrowheads) implanted into the mesenchyme. Beads contained BSA (control; F,G), Bmp4 (H,I) or noggin (J). The uneven white outline of the explant in J is from remnants of the filter on which the explant was cultured. (K-P) Inhibition of hedgehog signaling by cyclopamine (20 µM; n-p) reduces the expression of Foxf genes (L,M,O,P) to the same extent as of a known hedgehog target, Ptch1 (K,N). E12.5 intestinal explants were cultured for 24 hours in the presence or absence of cyclopamine and analyzed by whole-mount in situ hybridization.

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