ICANCKR RKSKARCH 56. 4Õ5R-436I. October 1. ]'»6]
Advances in Brief
Cell Cycle-dependent Expression of TAPI, TAP2, and HLA-B27 Messenger RNAs
in a Human Breast Cancer Cell Line1
Recep S. Alpan, Ming Zhang, and Arthur B. Pardee2
Divisions oÃ-Cell Growth (tiìdRegulation ¡K.S. A.. A. B. P.\ and Cancer Genetics ¡M.Z.¡.Dana-Furher Cancer Institute. De/hÃ-rtinent of Biological Chemistry inni Molecular
Pharmacology. Harvard Medical School. Boston Massachusetts 02115
Abstract
Tumor cells are generally poorly responsive to immunotherapy. The
results presented here suggest that antigen presentation of somatic tumor
cells may he diminished greatly in quiescence and may he determined in
part by growth regulation. Peptides produced by proteasomes are trans
ported into the endoplasmic reticulum by transporter proteins TAP-I and
TAP-2, where they bind and stabilize MHC class I molecules required for
antigenic presentation on the cell surface. TAP-1 and TAP-2 mRNAs were
undetectable in quiescent, serum-deprived human breast cancer cells
(2IPT). They appeared 10 h after serum induction, near the G,-S bound
ary. In contrast, III. \-l!27 mKNA was hiphasically up-regulated. These
mRNAs were significantly down-regulated in most tissues that contain
mainly terminally differentiated, nonprolif'erating cells. All of the inves
tigated breast cancer cell lines showed lower expression levels of these
mRNAs than did the corresponding normal cells.
Introduction
To identify growth-related genes in human breast cancer cells, we
recently compared mRNA expression by the method of differential
display ( 1, 2) in cells synchroni/.ed at quiescence (G„) and late G, (3).
From among several differentially expressed mRNAs. we identified
and cloned a cDNA that revealed 99% identity to human TAP-1
mRNA. MHC class I molecules present short peptides on the cell
surface for recognition by CD8+ CTLs (4). The short peptides are
processed by degradation of self- and foreign proteins in the cytosol
by a multisubunit protease called the proteasome (4-6). Short pep
tides produced by the proteasome are transported into the endoplasmic
reticulum by transporter proteins TAP-1 and TAP-2. which bind and
stabili/.e MHC class I molecules in the endoplasmic reticulum (4, 7).
Proteasome subunits LMP-2 and LMP-7 and peptide transporters
TAP-1 and TAP-2 are the products of genes that are located in the
class II region of chromosome 6 in humans. MHC class I heavy-chain
genes, including HLA-B27, are located in the MHC class I region in
the same chromosome (8). Inhibition of expression (9), inactivation of
TAP proteins (10), mutation (11). and proteasome inhibitor drugs (5.
11) were shown to result in insufficient presentation of antigenic
peptides with MHC class I molecules on the cell surface. This failure
of antigen presentation may result in escape of virus-infected and
transformed cells (viral or spontaneous) from immune recognition by
CTLs in vivo (12).
Proceeding from this finding, we now have analyzed TAP-1.
TAP-2, and HLA-B27 mRNA expression during the cell cycle. We
report here that the expression of these mRNAs is cell cycle depend
ent; they are down-regulated in quiescence created by serum depri
vation and are up-regulated in late G, following serum induction in
Received 6/17/96; accepted 8/15/96.
The costs of publication of this anide were defrayed in part by the payment of page
charges. This article must therefore be hereby marked advertisement in accordance with
18 U.S.C. Section 1734 solely to indicate this fact.
' This work was supported by N1H Grani GM-24571.
: To whom requests for reprints should he addressed, at Dana-Farher Cancer Institute.
44 Binney Street. Boston. MA 02115. Phone: (617) 632-3372; Fax: (617) 632-4680.
the human breast cancer cell line 21PT. Moreover, an average 3-fold
suppression in the expression levels of these mRNAs compared with
levels in normal breast epithelial cells was observed in a set of
investigated primary and metastatic breast cancer cell lines.
Materials and Methods
Cell Culture and Synchronization. The normal human mammary epithe
lial cell line X1N and human breast cancer cell lines 2IPT, 21NT. and 21MT2
used in this study were established as described (13) and kindly provided by
Ruth Sager (Harvard Medical School. Boston. MA). Other breast cancer cell
lines are from the American Type Culture Collection (Rockville. MD). To
create time course samples, 2IPT cells were starved in (i.5c/r I'etal bovine
serum for 85 h. At time 0. cells were released into complete medium contain
ing 10% fetal bovine serum, and samples were taken tor total RNA extraction
at indicated time points. Cell synchronization and cell cycle progression
following serum induction were monitored by flow cytometry as described
(14). For the lime course experiments. 2IPT cells were grown in «-MEM as
described (15). For growing normal mammary epithelial cells and breast
cancer cells. Dana-Farber Cancer Institute-l (DFCI-ll medium was used as
described (1ft).
cDNA Probes. TAP-1. TAP-2. and HLA-B27 full-length cDNAs were
kindly provided by Hidde Ploegh (Massachusetts Institute of Technology,
Cambridge. MA)
Total RNA Extraction and Northern Blot Hybridization. Total cellular
RNA extraction was performed with RNA/ol-B RNA extraction solution
(Biotecx Lab. Inc.. Houston. TX) according to the manufacturer's instructions.
Northern blot analyses were performed as described with minor variations
(17). Briefly. 20 /xg total RNA were resolved on a 1.1% agarose-1.7 M
formaldehyde gel and transferred to nylon membranes (MSI, Westboro. MA).
cDNAs used as probes were labeled with |'2P|dCTP by a random prime
labeling kit (Boehringer Mannheim Biochemicals. Indianapolis. IN) according
to the manufacturer's instructions, and hybridi/ations were performed as
described (17). For the analysis of mRNA expression in different tissues, a
human multiple-tissue Northern blot membrane (Clontech. Palo Alto, CA)
containing 2 jj.g polyadenylated RNA/lane was used. Northern blots were
quantitated by optical densitometric analysis (Bio-Rad GS-700 imaging den-
sitometer) and normalized with regard to ß-actinblots.
Results
TAP-1, TAP-2, and HLA-B27 mRNAs Are Up-Regulated fol
lowing Serum Induction in Human Breast Cancer Cells. Com
puter analysis of one of the cDNA clones identified by differential
display in late G, compared with quiescent cells showed 99% honiol-
ogy to TAP-1 mRNA in the GenBank (3). Northern blot analysis of
both the cDNA clone and the full-length TAP-1 cDNA on several
membranes gave identical size and expression patterns, confirming
the computer analysis. Next, we analy/ed the expression of TAP-2
and a class I heavy-chain (HLA-B27) mRNA during the cell cycle,
because they also are involved in the antigen presentation process on
the cell surface. Striking similarity of the expression patterns of
TAP-1 and TAP-2 was observed. Northern blots showed that both
TAP-1 and TAP-2 mRNAs were undetectable in the serum-deprived
cells. Both mRNAs were up-regulated in late G, after K) h of scrum
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Research.
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