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5.10. Additional RT-associated ribozymes
Structure-based searches for HDV-like ribozymes within the EST database,
known RT-containing mRNAs, and genomic repeat regions as defined by
RepeatMasker revealed many more ribozymes that map to various non-
LTR retrotransposons.
126
In addition to the R2 ribozymes in
Drosophila
,
the R2 elements of the termites
Kalotermes flavicollis
and
Reticulitermes
lucifugus
,
129
the sea squirt
Ciona intestinalis
,
130
the black-legged tick
Ixodes
scapularis
(GenBank
™
accession ABJB010506112), the horseshoe shrimp
Triops cancriformis
(GenBank
™
accession EU854578), and the Zebra finch
Taeniopygia guttata
131
were all found to harbor HDV-like ribozymes in their
5
0
terminal region. Other rDNA-associated retrotransposons with
ribozymes include the R4 element
132
in the human intestinal roundworm
Ascaris lumbricoides
, the horse intestinal roundworm
Parascaris equorum
, and
the R6Ag1 and R6Ag3 elements in
A
.
gambiae
.
133
With the exception of
the
A
.
gambiae
elements, all of these ribozymes possess the extended J1/2
region observed in the
Drosophila
isolates. Since this region can adopt
numerous conformations without abolishing self-cleavage, conservation
of the extended J1/2 region is likely attributable to each ribozyme's function
as an IRES. The JAM1 RTE
134
from the yellow fever mosquito
Aedes
aegypti
, the SARTPx2
133,135
element from the Asian swallowtail butterfly
Papilio xuthus
, and the Baggins retrotransposons from
Drosophila
are also
under the influence of HDV-like ribozymes. Both the JAM1 and SARTPx2
associated ribozymes, like the R6Ag1 and Ag3 ribozymes from
A
.
gambiae
,
lack the extended J1/2 region that is a hallmark of other rDNA-associated
retrotransposon ribozymes. These ribozymes do, however, contain P4 heli-
ces that are longer than those seen in the
Drosophila
R2 sequences, perhaps in
part to compensate for the loss of stability associated with a shorter J1/
2 motif.
Several ribozymes have been identified upstream of regions that display
high similarity to RT-coding motifs in a number of organisms, including
A
.
aegypti
, the butterfly
Heliconius numata
, the flatworm
Schistosoma mansoni
,
the sea slug
Aplysia californica
, the common sea urchin
Paracentrotus lividus
,
the Indonesian coelacanth
Latimeria menadoesnsis
, the sea lamprey
Petromyzon marinus
, the little skate
Leucoraja erinacea
, and the painted turtle
Chrysemys picta
.
5.11. HDV-like ribozyme cores similar to drz-Agam-1
In addition to drz-Agam-like ribozymes found in retrotransposons, a num-
ber of other sequences have been identified in genomes and ESTs from
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