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Gli proteins

Gli proteins mediate transcriptIonal control by Hedgehog signaling in vertebratesIn contrast to the fly, vertebrates have not one but three Ci-related proteins, designated Gli1, Gli2, and Gli3 (after glioblastoma), all of which have been implicated in mediating the activities of the various Hh proteins. The situatIon in vertebrates is further complicated by the fact that the genes encoding at least two of these, Gli1 and Gli3, are themselves transcriptIonal targets of Hh signaling: Gli1 is activated and Gli3 repressed in response to Hh signals (Marigo et al. 1996b; Lee et al. 1997). As usual, genetics has been a powerful tool in addressing the activities of these proteins, although as in the fly, the complexity of Gli functIon has complicated the interpretatIon of such analyses. Thus, for instance, although studies of mutant combinatIons in the mouse have shown genetic interactIons between Gli1 and Gli2 in the diencephalic regIon of the brain (Park and Bai 2000), Gli2 and Gli3 in skeletal development (Mo et al. 1997), Gli1 and Gli2, and Gli2 and Gli3 in lung development (Motoyama and Liu 1998; Park and Bai 2000), and Gli2 and Gli3 in tooth development (Hardcastle et al. 1998), the mechanistic actIon of the different Gli proteins in each of these organs remains unclear. Such mutant analyses have been complemented by studies in cell culture. InterpretatIon of these data, however, is also problematic, not least because in several cases, cells that are not responsive to Hh signals have been used, whereas in other instances, the experiments have not addressed the response of Gli proteins to the presence or absence of Hh ligand. Given the many levels of posttranscriptIonal regulatIon of Ci activity, in particular the Hh-dependent suppressIon of cleavage that alters the balance between repressor (cleaved) and activator (uncleaved) forms, conclusIons from these experiments should be treated with some cautIon. These studies have included the generatIon of modified forms of Gli

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