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Background  In  Drosophila melanogaster  the  Enhancer of split-Complex [E(spl)-C]  consists of seven highly related genes encoding basic helix-loop-helix (bHLH) repressors and intermingled, four genes that belong to the Bearded (Brd) family. Both gene classes are targets of the Notch signalling pathway. The  Achaete-Scute-Complex [AS-C]  comprises four genes encoding bHLH activators. The question arose how these complexes evolved with regard to gene number in the evolution of insects concentrating on Diptera and the Hymenoptera  Apis mellifera .    Results  In Drosophilids both gene complexes are highly conserved, spanning roughly 40 million years of evolution. However, in species more diverged like  Anopheles  or  Apis  we find dramatic differences. Here, the  E(spl)-C  consists of one bHLH ( mβ ) and one Brd family member ( mα ) in a head to head arrangement. Interestingly in  Apis  but not in  Anopheles , there are two more  E(spl)  bHLH like genes within 250 kb, which may reflect duplication events in the honeybee that occurred independently of that in Diptera. The  AS-C  may have arisen from a single  sc/l'sc  like gene which is well conserved in  Apis  and  Anopheles  and a second  ase  like gene that is highly diverged, however, located within 50 kb.    Conclusion   E(spl)-C  and  AS-C  presumably evolved by gene duplication to the nowadays complex composition in Drosophilids in order to govern the accurate expression patterns typical for these highly evolved insects. The ancestral ur-complexes, however, consisted most likely of just two genes:  E(spl)-C  contains one bHLH member of  mβ  type and one Brd family member of m α  type and  AS-C  contains one  sc/l'sc  and a highly diverged  ase  like gene.

The Enhancer of split and Achaete-Scute complexes of Drosophilids derived from simple ur-complexes preserved in mosquito and honeybee