Helitron Transposons on the Sex Chromosomes of the Platyfish Xiphophorus maculatus and Their Evolution in Animal Genomes

The sex-determining region of the sex chromosomes of the platyfish Xiphophorus maculatus contains several copies of a recently described class of DNA transposons called Helitrons, which probably transpose through a mechanism involving rolling circle replication. The unique open reading frame of platyfish elements encodes a 2816 amino-acid protein with helicase and replication initiator (Rep) domains, which are hallmarks of Helitrons. Like previously described elements from zebrafish but unlike sequences from plants, insects and nematodes, platyfish Helitrons also encode a C-terminal apurinic-apyrimidinic-like endonuclease probably captured from a non-longterminal- repeat retrotransposon. A cysteine protease domain related to the Drosophila ovarian tumor (OTU) protein was identified in the N-terminal part of the platyfish sequence. Putative endonuclease and protease have been acquired sequentially at least 600 million years ago and maintained functional in elements from sea urchin, lancelet and teleost fish, implying an important role for these domains in the transposition mechanism. Apparently intact Helitron elements are transcribed in Xiphophorus, and insertion polymorphism was observed between related fishes in different poeciliids. These observations suggest that Helitron transposons are still active in the genome of platyfish and related species, where they might play a role in the evolution of sex chromosomes and other genomic regions.