ELSEVIER Biochimica et Biophysica Acta 1218 (1994) 439-442
BB
Biochi~ic~a
et Biophysica A~ta
Short Sequence-Paper
Isolation and characterization of the tandemly repeated U6 genes
from the sea urchin Strongylocentrotus purpuratus
Sameer A. Sakallah 1, Devon R. Norton, Wei Zhang, William F. Marzluff *
Department of Biochemistry and Biophysics and Program in Molecular Biology and Biotechnology, University of North Carolina at Chapel Hill,
Chapel Hill, NC 27599, USA
Received 10 December 1993
Abstract
The tandemly repeated U6 genes were isolated from the sea urchin Strongylocentrotuspurpuratus. Each 1.8 kb repeat unit
contains a single U6 RNA sequence. There are no sequence similarities between the U6 promoter and other sea urchin snRNA
genes, other than a long polypyrimidine tract 3' of the U6 sequence.
Key words: Echinoderm; snRNA; U6 snRNA; (Sea urchin)
The sea urchin U1 and U2 snRNA genes expressed
in early sea urchin embryos are encoded in separate
tandemly repeated units [1,2]. The tandemly repeated
genes are expressed in oogenesis and early embryogen-
esis and then are silenced between the blastula and
gastrula stage [2-4]. All the spliceosomal snRNAs are
synthesized at a high rate from the 32-cell stage until
after hatching [5]. It is likely that each of the snRNAs
is encoded in a tandemly repeated gene set and that
these genes are coordinately regulated during embryo-
genesis. Here we report the isolation of one of the
tandemly repeated U6 snRNA genes from the sea
urchin Strongylocentrotus purpuratus.
Repeated screening of sea urchin genomic libraries,
which had yielded U1 [6,7] and U2 [2] snRNA genes
yielded only U6 pseudogenes, which contained trun-
cated U6 RNA sequences (S.A.S and W.F.M., unpub-
lished data). Since other sea urchin snRNAs are en-
coded by tandemly repeated genes and tandemly re-
peated genes may be under-represented in phage li-
* Corresponding author. Fax: + 1 (919) 9666821.
1 Present address: Department of Pathology, Division of Medical
Diagnostics, 1507W Biomedical Tower, University of Pittsburgh
Medical Center, Pittsburgh, PA 15261, USA.
The sequence data reported in this paper have been submitted to the
EMBL/GenBank Data Libraries under the accession numbers
X76389 and X76546.
0167-4781/94/$07.00 © 1994 Elsevier Science B.V. All rights reserved
SSDI 0167-4781(94)00038-5
braries, we cloned the U6 repeat using the polymerase
chain reaction. The sea urchin U6 RNA sequence was
determined by dideoxysequencing using reverse tran-
scriptase and a primer complementary to the conserved
U6 3' end. The sequence was confirmed by amplifying
U6 RNA by RT-PCR using primers complementary to
the 5' and 3' ends of the deduced U6 sequence (S.A.S.
and W.F.M., unpublished results). A repeat unit can be
isolated using two primers from the U6 coding region,
using the strategy shown in Fig. 1. The sequences of
the primers are given in the legend to Fig. 1. Primer 1
is nts 86 to 105 of the U6 sequence, and primer 2 is
complementary to nts 90 to 71 of the U6 sequence with
an EcoRI site added at the 5' end to facilitate cloning.
The amplified products were cloned into the plasmid
pUC18, and bacteria containing the U6 gene were
selected by colony hybridization using a 50 nt oligo-
nucleotide containing the central conserved region of
U6 snRNA as a probe. The complete sequence of the
1.82 kb insert (SpU6.1) was determined by dideoxyse-
quencing [8] using Sequenase (U.S. Biochemicals) or by
automated DNA sequencing. Since the amplified DNA
did not contain the complete U6 coding region, a 440
nt fragment containing the U6 coding region was am-
plified using primers located 5' and 3' of the coding
region (primers 3 and 4 in Fig. 1A). Primer 3 contained
nts -283 to -255 of the SpU6.1 repeat, including the
ClaI site at -265. Primer 4 was complementary to nts
+51 to + 32 and has an HindlII site added to facili-