Elements of the Drosophila bithorax complex that mediate repression by polycomb group products

The Polycomb (Pc) group genes encode repressors that restrict expression of homeotic genes to precise domains along the anterior-posterior (A-P) axis. We describe germline transformation constructs, containing portions of the bxd, iab-2, or iab-3 regulatory regions of the bithorax complex (BX-C), that are controlled by Pc group products in embryos. There are multiple BX-C elements that mediate Pc group control, which we call Pc group response elements (PREs), and they can work when removed from the normal BX-C context. These constructs are each regulated by the genes Polycomb (Pc), extra sex combs (esc), Enhancer of zeste (E[z]), polyhomeotic (ph), Polycomb-like (Pcl), Sex comb on midleg (Scm), Posterior sex combs (Psc), and Additional sex combs (Asx), consistent with multiple Pc group products acting together. Depending upon context, a PRE from the iab-3 region can restrict expression in different A-P positions. Thus, PREs are not specialized for particular parasegments, suggesting that Pc group products do not directly specify A-P boundaries of homeotic expression. Instead, the results support the idea that Pc group products provide stable memory or imprinting of boundaries which are initially specified by gap and pair-rule regulators.