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Fig. 7. Model for Gli2 and Gli3 function in somite myogenesis. Dorsoventral and mediolateral patterning of the somite is dictated in part by the opposing action of a ventral gradient of Hedgehog signalling (in red) and a lateral gradient of Bmp4 signalling (in blue). In the medial somite, the combined activity of Gli2A and Gli3A, converted from Gli3R into Gli3A by Shh, leads to the activation of Myf5 in DML cells and the activation of Pax1 in sclerotomal cells. Myf5 is also a target of Gli2A and Gli3A in the epaxial myotome, although a repressor function of Gli3 is revealed in Gli2–/–Gli3+/– embryos. In the lateral somite, characterised by low levels of Shh signalling and high levels of Bmp4 signalling, Myod1 and Lbx1 expression is controlled by Gli3R and Smad. We suggest that Gli3R could directly control Lbx1, Myf5 and Myod1 levels, and/or could interfere with Bmp4 signalling via an interaction with Smad in the control of Lbx1, Myf5 and Myod1 levels in the VLL and hypaxial myotome. Additional Gli/Shh-independent mechanisms of Myf5 activation are active in both the medial and lateral somite, possibly mediated by Wnt and/or FGF signalling.





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