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Prx1 and Prx2 are upstream regulators of sonic hedgehog and control cell proliferation during mandibular arch morphogenesis

Derk ten Berge1,*, Antje Brouwer1, Jeroen Korving1, Mark J. Reijnen1, Estia J. van Raaij1, Fons Verbeek1, William Gaffield2 and Frits Meijlink1,{ddagger}

1 Hubrecht Laboratorium, Netherlands Institute for Developmental Biology, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands
2 Western Regional Research Center, ARS, USDA, Albany, CA 94710, USA*Present address: Stanford University Medical Center, Beckman Center for Genetic and Molecular Medicine, 279 Campus Drive, Stanford, CA 94305-5323, USA



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  		Fig. 1. Fgf8 expression in the mandibular arch. (A,B) Fgf8 whole-mount in situ hybridization on dissected mandibular arch region of E11 wild-type (A) and Prx1-/-Prx2-/- (B) embryo, oral view. Dorsal side is upwards. Arrows in A indicate medial border of the Fgf8 expression domain in the wild type. In the mutant, the Fgf8 expression domain straddles the midline (arrow in B). Also visible in B is the distension at the midline (arrow), where the epithelium has detached from the mesenchyme. (C,D) Direct comparison of Fgf8 and Prx1 expression. Adjacent transversal sections through the mandibular arch area of E11 wild-type embryos were hybridized with radioactive Fgf8 and Prx1 probes. Double exposure of bright- and dark-field are shown; a red filter was used during darkfield photography. mand, mandibular arch mesenchyme; max, maxillary process mesenchyme.

 


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  		Fig. 2. Deformations and changes of patterning genes in mandibular arch of E10.5 embryos. (A) Prx2 expression in Prx2+/- embryo visualized by ß-galactosidase staining (blue). In situ hybridization with Dlx2 (B,C) and Alx3 (D,E) probes on nearby sections of wild-type (B,D) and Prx1-/-Prx2-/- (C,E) embryos. Expression is indicated in red. Owing to small differences in stage and orientation of embryos, sections at comparable levels do differ in shape and size. (F,G) The shift of the expression domains of Dlx2 and Alx3 in Prx1-/-Prx2-/- embryos (G) compared with wild-type embryos (F). Arrows in C,E indicate abnormal epithelium. Dorsal is upwards.

 


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  		Fig. 3. Detection of BrdU labeling in transverse sections of the mandibular arch of E10.5 wild-type (A,C) and Prx1-/-Prx2-/- mutant (G,I) embryos. (B,D,F,H) False color representations of A,C,G,I (respectively) to aid in interpretation (see Materials and Methods section). The strength of labeling is indicated by a color series ranging from yellow (weakest) to dark red (strongest), corresponding to ranges of arbitrary units as indicated. Boxed areas in the false color figures indicate areas in which the mutant shows reduced labeling compared with the wild type. White horizontal line in left lower corner of D indicates 48 pixels. (E,J) Nearby sections from wild-type (E) and Prx1-/-Prx2-/- (J) embryos showing expression of Shh in red. Dorsal is upwards.

 


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  		Fig. 4. Decrease of Shh expression in Prx mutants in time. Radioactive hybridization on cross-sections of embryos of various stages as indicated. (A,C,E) Wild type; (B,D,F) Prx1-/-Prx2-/- mutant.

 


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  		Fig. 5. Effects of jervine on lower jaw and limb development. (A) Exposure of embryos to jervine at E9.5 results in truncation of the distal lower jaw and loss of the incisors. Proximal is upwards. (B) Jervine exposure at E9.5 induces oligosyndactyly in the forelimb (left), but mainly carpal fusions in the hindlimb (right).

 


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  		Fig. 6. Reduction of lower jaw in Ptc transgenic embryos. E14.5 Ptc transgenic embryo (B) shows a shorter jaw than a wild-type example (A). (C,D) For comparison, skeletal staining of Prx mutant embryos are included to reveal similarities in mandibular phenotype. de, dentary; Me, Meckel’s cartilage.

 


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  		Fig. 7. Relationship between Prx1/Prx2, Shh and cell proliferation in the E10.5 mandibular arch. Prx1 and Prx2 in medial mandibular arch mesenchyme regulate the production of an unknown signaling factor (?). This factor signals to the oral epithelium and stimulates the expression of Shh. Shh signals back to the mesenchyme and stimulates cell proliferation.

 

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© The Company of Biologists Ltd 2001