Drosophila telomeres: an example of co-evolution with transposable elements.

Telomeres have a DNA component composed of repetitive sequences. In most eukaryotes these repeats are very similar in length and sequence and are maintained by a highly conserved specialized cellular enzyme, telomerase. Some exceptions of the telomerase mechanism exist in eukaryotes of which the most studied are concentrated in insects, and from these, Drosophila species stand out in particular. The alternative mechanism of telomere maintenance in Drosophila is based on targeted transposition of 3 very special non-LTR retrotransposons, HeT-A, TART and TAHRE. The fingerprint of the co-evolution between the Drosophila genome and the telomeric retrotransposons is visible in special features of both. In this chapter, we will review the main aspects of Drosophila telomeres and the telomere retrotransposons that explain how this alternative mechanism works, is regulated, and evolves. By going through the different aspects of this symbiotic relationship, we will try to unravel which have been the necessary changes at Drosophila telomeres in order to exert their telomeric function analogously to telomerase telomeres, and also which particularities have been maintained in order to preserve the retrotransposon personality of HeT-A, TART and TAHRE. Drosophila telomeres constitute a remarkable variant that reminds us how exceptions should be treasured in order to widen our knowledge in any particular biological mechanism.