Distal-less is a Downstream Gene of Deformed Required for Ventral Maxillary Identity

The homeotic selector (HOM) proteins are required for the diversification of the anterior-posterior axis of the Drosophila body plan, assigning unique identities to regional domains of cells comprising one or a few parasegments or segments. The HOM proteins apparently accomplish this task by the transcriptional regulation of numerous downstream genes. At present few downstream genes are known, so models of how downstream genes mediate HOM functions are based more on intuition than information. Our results indicate that Distal-less is a downstream gene of the HOM gene Deformed, and Distal-less function is required for the elaboration of a subset of the maxillary epidermal identities specified by Deformed. The regulatory effect of Deformed on Distal-less is mediated by a ventral maxillary-specific enhancer located 3' of the Distal-less transcription unit. We propose that Deformed and Distal-less, both of which encode homeodomain transcription factors that are persistently expressed in ventral maxillary cells, combinatorially specify a subsegmental code required for a group of cells to differentiate maxillary cirri.