[CANCER RESEARCH 52, 3931-3937. July 15. 1992]
Genetic Modification of a Murine Fibrosarcoma to Produce Interleukin 7
Stimulates Host Cell Infiltration and Tumor Immunity1
William H. McBride,2 J. Dean Thacker, Sian Coinora, James S. Economou, David Kelley, Donna Hogge,
Steven M. Dubinett, and Graeme J. Dougherty
Departments of Radiation Oncology /(*'. H. M., S. C.J, Surgery- ¡J.S. £./.Pulmonary and Critical Care Medicine fS. M. D., D. A'./, and the Jonsson Comprehensive
Cancer Center (H'. H. M., J. S. E., S. M. D.], University of California at Los Angeles School of Medicine, Los Angeles, California 90024; Terry Fox Laboratory
[J. D. T., D. H., G. J. D.], British Columbia Cancer Research Centre, Vancouver, British Columbia, V5ZIL3, Canada; and Departments of Medical Genetics
[J. D. T., D. H.l and Pathology ¡G. J. D./, University of British Columbia, Vancouver, British Columbia, l'6TIH'5, Canada
ABSTRACT
Retroviral-mediated gene transfer was used to introduce and express
the gene for murine interleukin 7 (II,-7) in a fibrosarcoma tumor (FSA).
The tumorigenicity of these genetically modified FSA cells was greatly
decreased in immunologically intact syngeneic mice but was unaltered in
T-cell-deprived mice. IL-7-infected tumors that did grow in intact ani
mals from large size inocula did so slowly and had a high incidence of
spontaneous regression. Furthermore, mice that had rejected tumors
became specifically immune to challenge with uninfected parental tumor
cells.
IL-7-infected FSA growing in intact mice were heavily infiltrated with
host T-cells that were presumably responsible for slow growth and
tumor regression, and tumor cells were in the minority. Fluorescence-
activated cell sorter analysis showed that there was a 530% increase in
T-cells in IL-7-infected FSA compared with control tumors. CD8+ T-
cells were particularly elevated, but CD4+ lymphocytes were also in
creased in number, as were eosinophils and basophils. The
CD4+:CD8+ ratio in IL-7-infected FSA was 1:1.7 in comparison to
1:0.6 in control tumors. Lymphocytes isolated from IL-7-producing tu
mors had greatly enhanced cytotoxicity towards uninfected, parental
FSA cells. Killing of non-cross-reacting fibrosarcoma line was also in
creased but to a much lesser extent.
Injection of recombinant human IL-7 directly into established FSA
tumors slowed their growth and, in a significant number of instances,
caused complete regression. Mice that had rejected tumor became spe
cifically immune. The dose that was needed for this effect was, however,
somewhat large: 20 *igtwice daily for 10 days. This result contrasts with
the efficacy of IL-7 gene infection in stimulating responses to the same
tumor. These considerations make IL-7 a good candidate for tumor-
directed cytokine gene therapy.
INTRODUCTION
The insertion of genes encoding various cytokines into tumor
cells has proven a useful approach with which to investigate the
molecular mechanisms that regulate in situ immune responses.
Murine tumor cells expressing genes for IL-2-1 (1-4), IL-4
(5, 6), IFN-7 (4, 7), and TNF-o (8, 9) have been found to have
decreased tumorigenicity and slower growth in vivo. IL-2 (3, 10)
and IFN-7 (7) were shown to mediate their effects through
mechanisms that were largely T-cell-dependent. In contrast,
tumor cells expressing IL-4 have been reported to induce re
sponses that were mainly T-cell-independent with contributions
Received 12/16/91; accepted 5/5/92.
The costs of publication of this article were defrayed in part by the payment of
page charges. This article must therefore be hereby marked advertisement in accor
dance with 18 U.S.C. Section 1734 solely to indicate this fact.
1This work was supported by grants from the National Cancer Institute, the
British Columbia Health Care Research Foundation, and the National Cancer
Institute of Canada. G. J. D. is a National Cancer Institute of Canada Career
Scientist.
2 To whom requests for reprints should be addressed, at Department of Radia
tion Oncology, UCLA School of Medicine. University of California, Los Angeles,
Los Angeles. CA 90024-1714.
3 The abbreviations used are: IL. interleukin; TNF-«,tumor necrosis factor a;
LAK, lymphokine-activated killer; TD?(). number of cells that had to be injected for
tumors to form in 50% of s.c. sites; IFN-7, Interferon 7; FCS. fetal calf serum.
from eosinophils and macrophages (5,6). TNF-a also appeared
to function by recruiting inflammatory cells, in particular those
bearing the complement receptor CR3 (8). Significantly, in
these studies mice that had eliminated the primary tumor fre
quently developed lasting tumor-specific immunity. These re
ports raise the possibility that tumor cells, genetically engi
neered to produce cytokines, may prove useful as immu-
notherapeutic agents. Such cells could deliver cytokines in a
prolonged fashion at high local concentrations without the side
effects that are frequently associated with the systemic delivery
of large bolus doses of these active biological agents. Moreover,
because the cytokines are delivered near tumor-reactive lym
phocytes, antitumor responses may be specifically enhanced.
IL-7 was originally described as a bone marrow stromal cell-
derived cytokine that stimulated the growth of pre-B-cells (11).
It was subsequently found also to function as a growth factor for
thymocytes (12, 13) and mature CD4+ and CD8+ T-cells (14,
15) and to stimulate resting T-cells to proliferate both directly
and through an IL-2-dependent pathway (14-19). This latter
activity can be attributed to IL-7-induced expression of IL-2
receptors. IL-7, therefore, acts to enhance T-cell competence
and lymphokine responsiveness in a manner similar to, but
independent of, IL-1 and IL-6 (15-18). IL-7 has also been
shown to help in the generation of cytotoxic T-cells (12, 20-22)
and LAK cells (21, 23, 24) and to stimulate the secretion of
IL-10, IL-6, and TNF-a by monocytes (25). Taken together,
these data suggest that IL-7 may play a role in the induction as
well as the expression of antitumor responses and to be a good
candidate for cytokine-mediated gene therapy. To test this pos
sibility, we have examined the immunological effects that result
from the introduction of the IL-7 gene into a murine fibrosar
coma of moderate immunogenicity.
MATERIALS AND METHODS
Mice. Experiments were performed on female C3Hf/Sed//Kam
mice aged 10-14 weeks. The animals were bred and maintained in our
defined-flora, specific pathogen-free colony at University of California-
Los Angeles. T-cell-deprived mice were prepared by thymectomy and
concurrent splenectomy at 4 weeks of age followed by lethal whole body
irradiation to 10.0 Gy and reconstitution with IO6 bone marrow cells
that had previously been treated with anti-Thyl.2 (monoclonal 30-H-
12; American Type Culture Collection, Rockville, MD) (26, 27). These
mice are immunocompromised and can support the growth of many
human tumor cell lines.
Tumor and Tumor Cell Lines. FSA is a fibrosarcoma that was in
duced in C3Hf/Sed//Kam mice by treatment with methylcholanthrene
(28). It is moderately immunogenic, grows progressively, and is well-
characterized with respect to the immunological responses it generates,
including its intratumoral host cell populations (26, 27, 29-31). We
have failed to find evidence for endogenously expressed retroviruses in
cultured FSA cell lines or antibody responses to viral antigens in mice
bearing FSA tumors (32). For cytokine gene transfer experiments, an in
vitro cell line was established from tumor that had been passed in vivo
3931
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