(CANCER RESEARCH 49,1422-1428, March 15,1989]
Expression of Anionic Glutathione-5-transferase and P-Glycoprotein Genes in
Human Tissues and Tumors
Jeffrey A. Moscow,1 Craig R. Fairchild, Mary Jane Madden, David T. Ransom, Harry S. Wieand, Erin E. O'Brien,
David G. Poplack, Jeffrey Cossman, Charles E. Myers, and Kenneth H. Cowan
Medicine Branch [J. A. M., C. R. F., M. J. M. E. E. O., C. E. M., K. H. C.], PediatrìeOncology Branch [D. G. P.], and Laboratory of Pathology fJ. C.J, Division of
Cancer Treatment, National Cancer Institute, Bethesda, Maryland 20892; and théDepartment of Medical Oncology [D. T. R., H. S. W.], Mayo Clinic, Rochester,
Minnesota 55905
ABSTRACT
The development of multidrug resistance in MCF-7 human breast
cancer cells and the acquisition of broad resistance to xenobiotics in rat
hyperplastic nodules are both associated with increased P-glycoprotein
(mar) gene expression as well as changes in activities of intracellular
(Intoxication enzymes; among these changes is a significant increase in
the activity of the anionic isozyme of glutathione-S-transferase (GST).
We have isolated a cDNA encoding the human anionic glutathione-5-
transferase, (ìS'1'ir-l, from a cDNA library constructed from multidrug-
resistant MCF-7 cells. The deduced amino acid sequence of GSTr-1
shows that while the human anionic GST displays 85% nucleotide and
amino acid sequence homology to the rat anionic isozyme, it is markedly
less related to human basic GST isozymes. We have examined the
expression of GST* and P-glycoprotein in 170 specimens of human
tissues and tumors. P-Glycoprotein RNA expression was positive in eight
of 23 lymphomas and two of 12 colon tumors; however, many other
normal and malignant tissues, including lung, bladder, and breast tumors,
had low or undetectable levels of P-glycoprotein RNA expression. In
contrast, GST»was readily detected in a wide variety of normal and
malignant tissues. The level of GSTr mRNA expression in normal tissues
was heterogeneous, with lowest levels found in liver and the highest
levels found in lung, esophagus, and placenta. GSTr was also variably
expressed in human tumors, with the lowest relative levels occurring in
lymphoma and breast cancer and the highest levels found in lung cancer
and head and neck tumors. In addition, comparison of paired specimens
from the same patient indicated that GST n-expression was increased in
many tumors relative to matched normal tissue.
INTRODUCTION
The phenomenon of MDR,2 the ability of malignant cells in
vitro to develop cross-resistance to a broad range of structurally
dissimilar toxins, has been associated with decreased intracel
lular drug accumulation and with overexpression of a high-
molecular weight (A/r 130,000-180,000) membrane-associated
glycoprotein (P-glycoprotein) (1-5). Recent studies have shown
that MDR can be conferred upon drug-sensitive cells by transfer
of genes encoding P-glycoprotein (6-8). However, despite re
ports of increased P-glycoprotein expression in some human
tumors in the past few years (9-12), the role of P-glycoprotein
in clinical drug resistance is not yet clear.
We have recently reported the development of MDR in a
human breast cancer cell line selected for resistance to Adria-
mycin (Adr* MCF-7) (13). MDR in this cell line is associated
with a number of phenotypic and biochemical changes that are
remarkably similar to those reported to occur in vivo in rat liver
HNs (14), a well-characterized model of chemical carcinogen-
esis in which the transformed hepatocytes demonstrate broad-
spectrum cross-resistance to hepatic toxins (15). Both Adr*
Received 7/7/88; revised 11/18/88; accepted 11/29/88.
The costs of publication of this article were defrayed in part by the payment
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1To whom correspondence should be addressed at Building 10, Room 12C112,
National Cancer Institute, National Institutes of Health, Bethesda, MD 20892.
1The abbreviations used are: MDR, multidrug resistance; HNs, hyperplastic
nodules; GST, glutathione-S-transferase.
MCF-7 cells and rat HNs demonstrate decreased toxin accu
mulation accompanied by overexpression of the P-glycoprotein
gene as a manifestation of the MDR phenotype (16, 17). In
addition, however, broad spectrum resistance in both models is
associated with similar changes in the regulation of intracellular
detoxication reactions, including marked increases in glutathi-
one-S-transferase activity (13, 18-20).
The glutathione-S-transferases are a family of enzymes which
are involved in the metabolism of a broad range of xenobiotics
(21). These enzymes exist as homodimers or heterodimers. The
cytosolic forms of these enzymes are the products of three
separate gene families which are distinguished on the basis of
distinctive isoelectric focussing properties; basic, neutral, and
anionic. GST isozymes have in common their ability to detoxify
electrophilic and Hydrophobie substances with reduced gluta-
thione. In addition, GST isozymes have been shown to bind
covalenti) and noncovalently to toxins and to detoxify organic
peroxides to less reactive alcohols (21). Isozymes composed of
subunits from these different groups have different substrate
specificities and are immunologically distinct.
Recent studies have shown that the activity of the anionic
GST isozyme is increased in both rat HNs and Adr* MCF-7
cells. The rat anionic GST, GST-P, an isozyme usually present
only in low levels in liver, is markedly elevated in xenobiotic-
resistant rat HNs (18, 19). In addition, our laboratory has
reported that the human form of the anionic GST, GST*-, is
increased in multidrug-resistant Adr* MCF-7 cells when com
pared with the parental cell line (WT MCF-7) (13, 20).
In order to further explore the association between GSTir
and drug-resistance, and to determine whether GSTTr could be
a useful marker for clinical drug resistance, we have cloned the
cDNA encoding this enzyme and used this cDNA as a probe to
examine GSTjr expression in human tissues and tumors. We
also recently reported isolation of a human P-glycoprotein
sequence cDNA from Adr* MCF-7 cells (22) and have exam
ined the same specimens for P-glycoprotein expression in order
to compare the expression of these two genes in a variety of
human tumors.
MATERIALS AND METHODS
Nucleotide Sequence Analysis. GST«is a 725-base pair cDNA cloned
from a cDNA library constructed from RNA isolated from multidrug-
resistant Adr" MCF-7 cells and which was screened by hybridization
with a radiolabeled rat anionic GST probe, pGP5 (23), which was
kindly provided by M. Muramatsu. GSTi-l fragments were subcloned
into an M 13 vector (Bethesda Research Laboratories) and sequenced
using the dideoxy method with 3sS-labeled adenosine triphosphates.
Standard M-13 primers were used to sequence from the ends of the
fragments (Bethesda Research Laboratories) according to the manufac
turer's instructions. Additional oligonucleotide primers were con
structed to sequence areas from within the fragments (Midland Certified
Reagent Company, Midland, TX).
Hybridization Studies. Tumors and normal tissues for RNA analysis
from surgical pathological specimens were quick-frozen in liquid nitro
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