[CANCER RESEARCH54, 4851—4854, September 15, 1994]
Advances in Brief
Bcl-2 Inhibits T-Cell-mediated Cytolysis of a Leukemia Cell Line1
Toshihiko Torigoe, Juan A. Milan, Shinichi Takayama, Russell Taichman,2 Toshiyuki Miyashita,
and John C. Reed3'4
Oncogene & Tumor Suppressor Gene Program, La Jolla Cancer Research Foundation, La Jolla. California 92037
cells against antibody-based responses, since it does not prevent
complement-mediated cell death (10) and does not prevent phagocy
tosis (1 1), as might occur, for example, when tumor cells are coated
with antibody. The potential impact that Bcl-2 might have on cell
mediated immune responses, however, is more difficult to predict
because of the multiple mechanisms that CTLs, NK cells, LAK cells,
and tumor infiltrating lymphocytes have at their disposal for inducing
target cell death (reviewed in Ref. 12) and because the predominant
mechanism used appears to vary among different populations or
clones of T-, NK, LAK, and tumor-infiltrating lymphocyte cells. In
this regard, cell-mediated cytolysis can occur through mechanisms
consistent with either apoptosis or necrosis, depending on the partic
ular immune cell effectors and the target cells studied (reviewed in
Ref. 13). For the most part, C'TLs, NK, and related cells kill tumor
targets through two separate processes: (a) secretion of cytotoxic
granules which contain a number of potentially lethal molecules
including perform (a protein that resembles the C9 component of
complement and that pokes holes in membranes), AlT (which can
stimulate apopotosis in some types of cells probably via purinergic
receptors), proteases (that trigger DNA fragmentation and apoptosis
through a pathway which may require cdk kinases), and TIA-1 (an
RNA-binding protein that induces apoptosis through poorly under
stood mechanisms); and (b) expression of genes encoding cytotoxic
cytokines, such as TNFc-a, lymphotoxins, and the Fas ligand (14—
18). With regards to the latter, gene transfer-mediated elevations in
Bcl-2 protein levels have been shown to provide at least partial
protection against apoptotic cell death induced by TNF-a and anti
bodies to Fas (19, 20), suggesting that Bcl-2 could potentially render
tumor cells more resistant to CTL-induced killing in scenarios where
local secretion of TNF-a or expression of transmembrane forms of
TNF-a or Fas ligand play a predominant role in the cytolytic mech
anism. On the other hand, if perform-dependent mechanisms involv
ing loss of osmotic equilibrium and necrotic cell death predominated,
Bcl-2 would not be expected to provide resistance, by analogy to the
lack of protection seen previously for complement-mediated lysis
(10). In fact, it has been reported by others that Bcl-2 does not protect
against cytolysis induced by at least some allospecific CFLs (10, 21).
We show here, for the first time, however, that gene transfer-mediated
elevations in Bcl-2 protein levels can render a human leukemia cell
line relatively more resistant to DNA fragmentation and cytolysis
induced by a cloned T-cell that induces cell death through mecha
nisms consistent with apoptosis.
Abstract
The bcl-2 gene becomes dysregulatedin its expressionin a wide variety of
human cancers and has been shown to block both spontaneous and drug
induced cell death, thus conferring a selective survival advantage on malig
smut cells. The biochemical mechanism by which bcl-2 promotes cell survival
remains enigmatic but appears to involve a downstream event in an evolu
tionarily conserved cell death pathway. Here we report that gene transfer
mediated increases in Bd-2 protein levels in the human leukemia line Jurkat
render these cells more resistant to induction of DNA fragmentation and
cytolysis by a cloned T-celL The kiffing mechanism used by these particular
T-cells was consistent with apoptosis, as opposed to necrosis, in that DNA
degradation occurred as a prelysis event. The findings raise the possibffity
that dysregulation ofbcl-2 gene expression could play a role in the avoidance
of immune surveillancemechanisms by cancer cells.
Introduction
The bcl-2 gene was first discovered because of its involvement in
the t(14;18) chromosomal translocations that frequently occur in
non-Hodgkin's B-cell lymphomas and which move the bcl-2 gene
from its normal location at 18q21 into a cis-configuration with strong
enhancer elements associated with the immunoglobulin heavy-chain
locus at 14q32 (1). High levels of Bcl-2 protein production have also
been reported in a wide variety of human solid tumors and leukemias
in the absence of translocations or other gross alterations in the
structure of the bcl-2 gene, including adenocarcinomas of the prostate
and colon, small cell and non-small cell carcinomas of the lung,
nasopharyngeal carcinomas, neuroblastomas, acute myelogenous leu
kemias, and chronic lymphocytic leukemias (for examples, see Refs.
2—6).The Bcl-2 protein has been shown to contribute to neoplastic
cell expansion by blocking the normal physiological turnover of cells
that occurs due to programmed cell death (reviewed in Ref. 7). In
addition, gene transfer-mediated elevations in Bcl-2 protein levels
have been shown to render leukemia and tumor cells relatively more
resistant to induction of apoptosis by multiple types of chemothera
peutic drugs (8, 9). Bcl-2, therefore, may play a significant role, not
only in the origins of cancer, but also in its treatment.
Yet another aspect of tumor biology that determines the potential
for individual neoplastic clones to gain a selective growth advantage
in vivo concerns the issue of the immune surveillance mechanisms
that help to eradicate or keep in check abnormal cells in the body. In
this regard, both humoral (antibody-mediated) and cellular responses
can play a role in host defenses against cancer. Overproduction of the
Bcl-2 protein would probably not be expected to protect malignant
Received 6/20/94; accepted 8/1/94.
The cost of publication of this article 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.
I This study was supported in part by American Cancer Society Grant IM-708 and NIH
Grant CA-54957.
2 Present address: Department of Periodontics, School of Dentistry, University of
Michigan, Ann Arbor, MI 48109.
3 To whom requests for reprints should be addressed, at Oncogene & Tumor Suppres
tar Gene Program, La Jolla Cancer Research Foundation, 10901 North Torrey Pines Road,
La Jolla, CA 92037.
4 Scholar of the Leukemia Society of America.
Cells and Cell Culture. The cytolytic properties, immunophenotype, and
maintenance in culture of the particular clone of IL-2-dependent CFLL-2
cytolytic T-cells and various transfectants thereof have been described in detail
previously (22). CTLL-N-LCK and CT'LL-A-LCK cells represent transfectants
that received a G418-resistant plasmid in combination with expression plas
mids producing either normal p56-Lck or an “activated― version of Lck that
contains a tyrosine—'phenylalanine substitution at position 505.
Jurkat-NEO and Jurkat-BCL-2 cells are polyclonal populations of 0418-
resistant Jurkat T-cell leukemia cells that were subjected to electroporation
4851
Materials and Methods
Research.
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