Overlapping transcription units in Drosophila: sequence and structure of the Cs gene

The Cs gene lies between the functionally and evolutionarily related dopa decarboxylase (Ddc) and l(2)amd loci of Drosophila. The Cs and Ddc genes overlap at their 3' ends, implying that the transcription termination signals of these genes are polar, since each gene's primary transcript contains the complement of the other gene's transcription termination signals. The mature transcripts of the Cs and Ddc genes are complementary for a short distance and the primary transcripts may be complementary over thousands of base pairs. Despite intensive mutagenesis in this region, no mutations affecting the Cs transcript have been recovered although over 90 alleles of the two flanking genes (Ddc and l(2)amd) have been identified. Unlike the flanking Ddc and l(2)amd genes, the structure of the Cs gene and the temporal and tissue specificity of Cs expression are inconsistent with any structural or functional relatedness to the Ddc gene family. The internal structure of the Cs transcript is unlike that of most protein coding genes; it contains several open reading frames which are not situated favorably for efficient translation of the Cs message. This unusual internal structure may be the basis of the observed mutational silence of the Cs locus.