Homoeosis in Drosophila: a description of the Polycomb lethal syndrome

Adults heterozygous for dominant mutations at the haploabnormal Polycomb (Pc) locus display many homoeotic transformations. E. B. Lewis (1978) first described the cuticular morphology of lethal embryos homozygous for Polycomb mutant alleles, and suggested that the Pc+ gene product acts as a repressor of genes in the Bithorax gene complex. In the present work, we have further examined the Polycomb lethal syndrome by phase-contrast and scanning electron microscopy of whole mounts, and show that the phenotype is more complex than hitherto realized. Many cuticular features characteristic of more anterior body segments are partially or completely transformed to resemble those of more caudal segments; the posteriormost homoeotic features which develop are those of the 8th abdominal segment (rather than the 9th or 10th). Involution and dorsal closure of the head are usually incomplete, and the labium and dorsal head regularly develop cuticular features normally characteristic of abdominal segments. Not all homoeotic alterations are caudal, however, and embryos also display partial transformations of the 9th (or 10th) abdominal segment to 8th and of meso- and metathorax to prothorax. It should be emphasized that Polycomb mutations do not homoeotically transform segments as a whole; various cuticular markers differ considerably in their relative probability and extent of transformation, and a single segment may display features normally specific to several different segments. We suggest that these mutations result in an instability in the transmission of determined states.