The Gene Network That Regulates Head Organizer Activity in Is Differentially Regulated in Epidermis and Gastrodermis

head formation depends on an organizing center in which Wnt/β-catenin signaling, that plays an inductive role, positively regulates and , with Sp5 limiting expression and Zic4 triggering tentacle formation. Using transgenic lines in which the promoter drives eGFP expression in either the epidermis or gastrodermis, we show that promoter activity is differentially regulated in each epithelial layer. In intact animals, epidermal GFP activity is strong apically and weak along the body column, while in the gastrodermis, it is maximal in the tentacle ring region and maintained at a high level along the upper body column. During apical regeneration, :GFP is activated early in the gastrodermis and later in the epidermis. Alsterpaullone treatment induces a shift in apical expression towards the body column where it forms transient circular figures in the epidermis. Upon (RNAi), GFP activity is down-regulated in the epidermis while bud-like structures expressing GFP in the gastrodermis develop. (RNAi) reveals a negative autoregulation in the epidermis, but not in the gastrodermis. These differential regulations in the epidermis and gastrodermis highlight the distinct architectures of the network in the hypostome, tentacle base and body column of intact animals, as well as in the buds and apical and basal regenerating tips.