A model for the origin of internal eliminated segments (IESs) and gene rearrangement in stichotrichous ciliates

Abstract

A characteristic feature of ciliates (ciliated protozoans) is their nuclear dimorphism: the presence of two kinds of functionally different nuclei in the same cell—a micronucleus (MIC) and the macronucleus (MAC). In the stichotrichous group of ciliates the organization of DNA in the MIC is dramatically different from that in the MAC. Genes in the MIC consist of the sequence of segments, called MDSs, which are separated by short noncoding pieces of DNA, called IESs. Moreover, the order of MDSs in the MIC may be scrambled compared to their order in the MAC, and also some MDSs may be inverted with respect to each other. In this paper, we consider the evolutionary origin of this bizarre form of MIC genes, and in particular we postulate that the insertion of IESs as well as possible scramblings/inversions have resulted from a repair of one or more breaks in a DNA molecule. We propose a specific repair scheme, and postulate that this repair scheme applied to a coiled structure of a DNA molecule that has undergone multiple breaks can produce IES insertions and/or scrambled/inverted MIC gene patterns. All experimentally demonstrated as well as theoretical MIC gene patterns can be produced in this way.