Exon shuffling and other ways of module exchange

Thanks to recent improvements in techniques used for the detection of homologies, it is now clear that module exchange played a major role in protein evolution. Analysis of the genes of various modular proteins has identified a large number of cases where gene assembly was facilitated by intronic recombination - i.e., the proteins were formed by exon shuffling. Studies of the principles and mechanistic details of exon shuffling, however, revealed that this powerful evolutionary mechanism could become significant only after the appearance of spliceosomal introns typical of higher eukaryotes. Although exon shuffling is the most efficient way of constructing modular proteins, recent studies on the evolution of multidomain proteins of prokaryotes emphasize that intronic recombination is not an absolute prerequisite of module exchange.