One of the functions of telomeres is to counteract the terminal nucleotide loss associated with DNA replication. While the vast majority of eukaryotic organisms maintain their chromosome ends via telomerase, an enzyme system that generates short, tandem repeats on the ends of chromosomes, other mechanisms such as the transposition of retrotransposons or recombination can also be used in some species. Chromosome end regression and extension were studied in a medically important mosquito, the malaria vector Anopheles gambiae, to determine how this dipteran insect maintains its chromosome ends. The insertion of a transgenic pUChsneo plasmid at the left end of chromosome 2 provided a unique marker for measuring the dynamics of the 2L telomere over a period of about 3 years. The terminal length was relatively uniform in the 1993 population with the chromosomes ending within the white gene sequence of the inserted transgene. Cloned terminal chromosome fragments did not end in short repeat sequences that could have been synthesized by telomerase. By late 1995, the chromosome ends had become heterogeneous: some had further shortened while other chromosomes had been elongated by regenerating part of the integrated pUChsneo plasmid. A model is presented for extension of the 2L chromosome by recombination between homologous 2L chromosome ends by using the partial plasmid duplication generated during its original integration. It is postulated that this mechanism is also important in wild-type telomere elongation.Telomeres are essential chromosomal structures whose functional integrity is linked to cell cycle progression. Considerable efforts are being made to understand telomere structure and to target elongation mechanisms in order to develop new means of proliferation control (12). Every eukaryotic organism must compensate for terminal loss of DNA from chromosome ends because DNA polymerases cannot completely replicate the ends of linear chromosomes. So far, evidence for three different terminal elongation mechanisms has been established. Most eukaryotes have a short, tandemly repeated DNA sequence motif on their chromosome ends. These telomeric tandem arrays are extended by a specific reverse transcriptase, telomerase, which carries an internal RNA template (12). Alternatively, Drosophila melanogaster elongates its telomeres by a very different mechanism that is based on transpositions of specific retrotransposons, HeT-A and TART, to chromosome termini (22). In addition, recombination of repetitive telomeric ends has been considered as a possible elongation mechanism (4, 10, 42). This pathway has been well documented in yeast where telomeres are extended by telomerase, but recombination can be used as an efficient bypass mechanism for elongation (19,20,30,39).Understanding the mechanism of telomere elongation in the malaria-transmitting mosquito Anopheles gambiae may be useful for developing new strategies for vector control. Therefore, we examined the dynamics of chromosome length variation at the 2L chromosome end. For...