mRNA maturation in Trypanosoma brucei
depends upon trans-splicing, and variations in trans-splicing
efficiency could be an important step in controlling the levels of
individual mRNAs. RNA splicing requires specific sequence elements
including conserved 5´ splice sites, branch points,
pyrimidine-rich regions (poly(Y) tracts), 3´ splice sites
(3´SS) and, sometimes, enhancer elements. To analyze sequence
requirements for efficient trans-splicing, in the poly(Y) tract and
around the 3´SS, we constructed a luciferase-b-galactosidase
double-reporter system. By testing ~90 sequences, we demonstrated
that the optimum poly(Y) tract length is ~25 nt. Interspersing a
purely uridine-containing poly(Y) tract with cytidine resulted in
increased trans-splicing efficiency, whereas purines led to a large
decrease. The position of the poly(Y) tract relative to the 3´SS
is important: an AC dinucleotide at position -3 and -4 can lead to a
20-fold decrease in trans-splicing. However, efficient trans-splicing
can be restored by inserting a second AG dinucleotide downstream,
which does not function as splice site but may aid in recruitment of
the splicing machinery. These findings should assist the development
of improved algorithms for computationally identifying 3´SS, and
help to discriminate non-coding open reading frames from true genes,
in current efforts to annotate the T. brucei genome.