Schematic illustrations of the Tetrahymena telomerase TERT and TER subunits. (A) A linear map of the Tetrahymena telomerase major catalytic protein TERT, which consists of 1117 amino acids. The four domains: TEN, TRBD, RT (reverse transciptase domain) and CTE (carboxy-terminal element) are indicated. Also indicated are motifs that are conserved in all DNA polymerases and reverse transcriptases (1,2,3,A,B′,C,D,E), and other motifs that are only conserved in telomerases (IFD,T,QFP,CP,CP2,GQ). Of the three amino acids in the TEN domain, which are indicated in the drawing, W187 and Q168 were found by site-directed UV cross-linking and kinetic measurements to contact growing DNA chains in active core telomerase complexes (24). L14 was found to be specifically required for processive extension of telomeric DNA chains by the core telomerase (26). (B) Sequence and secondary structure of the Tetrahymena TER, which consists of 159 nucleotide residues. The template sequence and the regions TBE and TRE (TERT recognition element) are colored blue, violet and red, respectively.