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Journal of General Virology (1995), 76, 139 145. Printed in Great Britain 139
Sequence polymorphism in the Epstein-Barr virus latent membrane
protein (LMP)-2 gene
Pierre Busson, 1,~ Rachel Hood Edwards, 1 Thomas Tursz 2 and Nancy Raab-Traub ~*
1Department of Microbiology and Immunology and the Lineberger Cancer Research Center, University of North
Carolina, Chapel Hill, NC 27599-7295, USA and 2Laboratoire de Biologie des Tumeurs Humaines,
Institut Gustave Roussy, Villejuif, France
Latent membrane protein 2A (LMP-2A) is expressed in
Epstein-Barr virus transformed B lymphocytes in vitro
and has been detected in various types of EBV-
associated malignancies. LMP-2A interferes with mem-
brane signal transduction through phosphorylation of its
hydrophilic N-terminal domain and binding of the
cellular tyrosine kinases encoded byfvn and lyn. In vitro,
the domain can block calcium influx and participate in
signal transduction inducing cytokine production. These
two activities are differently affected by site-directed
mutagenesis of potentially phosphorylated amino acid
residues. Several potential tyrosine protein kinase rec-
ognition motifs have been identified including an antigen
recognition motif (ARAM). ARAMs are activated by
tyrosine phosphorylation that enables binding of tyro-
sine protein kinases such as lyn and fvn. To assess the
importance of potential sequence variation in natural
EBV infection and in tumourigenesis, the sequence of
the LMP-2A N-terminal domain was determined in 28
EBV isolates, including 14 fresh tumour isolates.
Comparison of the corresponding sequences with the
prototype B95 strain indicates that LMP-2 is generally
conserved with a few base pair changes resulting in
conservative amino acid changes in an occasional
isolate. However, five single-base loci were frequently
mutated, resulting in three patterns of sequence poly-
morphism in exon 1 of LMP-2A. The patterns did not
segregate with EBV Type 1 or Type 2 and were detected
in both lymphoid and epithelial tissues. Four of the most
frequent mutations at loci 166627, 166750, 166796 and
166805 (codons 23, 63, 79 and 82) could potentially
affect tyrosine protein kinase binding motifs. The pivotal
tyrosines (codons 74 and 85) and leucines (codons 77
and 88) of the LMP-2 ARAM were not affected in any
of the isolates, suggesting that ARAM function is
important for EBV infection in vivo. However, the inter-
spacing positions 79 and 82 were distinct in more than
50% of the isolates. These prevalent polymorphisms
could influence interaction of the LMP-2 cytoplasmic
domain with specific cellular ligand proteins.
Epstein-Barr virus (EBV) encodes two integral mem-
brane proteins, latent membrane proteins (LMPs) 1 and
2 (Kieff & Liebowitz, 1990). There are two forms of
LMP-2, LMP-2A and -2B, that are encoded by two
transcripts which are spliced across the terminal repeats
of the EBV DNA (Laux et al., 1988; Sample et al., 1989).
Therefore, their transcription requires episomal forms of
the viral genome or rare integrative events (Hurley et al.,
1991). Each transcript results from the alternative
splicing of a unique 5' exon 1, located at the right-end of
the genome, while exons 2 to 9, located at the left-end of
the genome, are common to both species. The LMP-2A
5' first exon (368 bp) encodes an N-terminal hydrophilic,
intracytoplasmic domain (119 aa) while the LMP-2B
first exon (154bp) is non-coding. The eight exons
common to LMP-2A and -2B encode a highly hydro-
* Author for correspondence. Fax +19199663015. e-mail
nrt@med.unc.edu
phobic transmembrane domain (1140 coding nucleo-
tides; 378 aa) (Kieff & Liebowitz, 1990; Rabson et al.,
1982).
The LMP-2A and -2B proteins are known to be
expressed in lymphoblastoid and Burkitt's lymphoma
cell lines (Frech et al., 1990; Longnecker & Kieff, 1990).
The LMP-2 genes are consistently transcribed in naso-
pharyngeal carcinomas (NPC) and LMP-2A transcripts
have also been detected in EBV-positive uncultured
peripheral blood lymphocytes (Brooks et al., 1992;
Busson et al., 1992; Qu & Rowe, 1992). The sequence of
two overlapping cDNAs representative of the entire
LMP-2A coding sequence was determined in C15, an
NPC established in nude mice (Busson et al., 1988, 1992).
When compared with the B95 sequence, exons 2 to 9
were highly conserved; on a segment of 1140 bp, five
mutations were detected with only one resulting in a
significant change, at codon 126, with the replacement of
a leucine by a proline. In contrast, nine mutations were
0001-2626 © 1995SGM