[CANCER RESEARCH 58. 1225-1230. March 15. 1998|
Interleukin 12 Gene Therapy of MHC-negative Murine Melanoma Métastases1
Patrizia Nanni,2 liarÃ-aRossi, Carla De Giovanni, Lorena Landuzzi, Giordano Nicoletti, Antonella Stoppacciaro,
Marii-Ila Parenza, Mario P. Colombo, and Pier-Luigi Lollini
Institute for Cancer Research, University of Bologna, 1-40126 Bologna [P. N., I. /?., C. D. G., P- L. L.]: National Cancer Institute of Genoa. Biolechnoh>x\ Satellite Unit of
Bologna, 1-40126 Bologna ¡L.L. G. N.]: Department of Experimental Medicine, University of Rome, 1-()Õ)I61Rome ¡A.S.l; and Experimental Oncologv [). Natitmal Cancer
Institute, 1-20133 Milan ¡M.P., M. P. C.]: Italy
ABSTRACT
Immunological gene therapy of cancer relies heavily on the activation of
T cells, but tumors with defects in MHC gene expression are not recog
nized by MHC-restricted T cells. To investigate the potential of cytokine
genes for the therapy of MHC-negative tumors, we transduced B78H1, a
class I-negative murine melanoma clone, with a polycistronic vector car
rying murine interleukin (II.I-I2 genes. The clones studied produced
400—25,000 pg/ml 11-12 their in vitro growth properties were similar to
those of parental cells. A complete inhibition of growth was observed in
vivo both after s.c. and i.v. administration of all II.-12 clones. II ,-12-
transduced cells were also used as a therapeutic vaccine in mice bearing
micrometastases by nontransduced parental cells. A significant (80-90%)
reduction in the number of lung nodules was obtained. Immunohisto-
chemical analysis and studies in immunocompromised hosts showed that
T cells and natural killer cells had a significant role in the elimination of
IL-12-releasing cells. In situ hybridization with cytokine probes detected a
strong increase in the proportion of leukocytes positive for IFN-y, tumor
necrosis factor a, H.-l/i. and IFN-inducible protein 10 at the site of
rejection of IL-12-engineered tumor cells. However, it was clear that the
loss of in vivo growth was also due to T-cell- and natural killer cell-
independent factors, possibly related to the antiangiogenic properties of
11-12. In conclusion, tumor therapy based on 11-12 gene transduction was
effective on a MHC-negative metastatic tumor, suggesting a possible
application to MHC-defective human neoplasms.
INTRODUCTION
The use of recombinant cytokines and of cells transduced with
cytokine genes, in association with the cloning of specific tumor
rejection antigens, has considerably increased the possibilities to
induce or to boost an antitumor immune response based mainly on T
lymphocytes. On the other hand, detailed studies of MHC gene
products revealed that defects in class I MHC expression are wide
spread among human tumors. In fact, according to some estimates,
most human tumors fail to express one or more class I MHC glyco-
proteins on their membrane (1, 2).
A causal link between specific MHC class I loss and immunoselec-
tion by T lymphocytes has not yet been established; however, it is
certain that the absence of a given MHC gene product prevents the
recognition by T lymphocytes of the antigenic peptides for which the
MHC product represents the restriction element (1).
The high prevalence of tumors with class I MHC defects indicates
that specific strategies should be devised to cope with tumors that
cannot be efficiently killed by MHC-restricted CTLs. A good candi
date for this type of application is IL-12,3 which is able to activate
both specific TH1 responses and nonrestricted NK effectors (3).
Received 10/15/97; accepted 1/16/98.
The costs of publication of this article were defrayed in part by the payment of page
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18 U.S.C. Section 1734 solely to indicate this fact.
1 Supported by grants from the Italian Association for Cancer Research, the Italian
Ministry for University and Research, and the National Research Council. I. R. is the
recipient of an Italian Association for Cancer Research fellowship.
2 To whom requests for reprints should be addressed, at Istituto di Cancerologia. Viale
Filopanti 22.1-40126 Bologna, Italy. Fax: 39-51-242-169; E-mail: nanni@cancer.unibo.it.
3 The abbreviations used are: IL. interleukin; rIL. recombinant IL: IP10. IFN-inducible
protein 10; TNF. tumor necrosis factor; NK. natural killer: niAb. monoclonal antibody.
IL-12 is a heterodimeric cytokine produced by antigen-presenting
cells, phagocytes, and granulocytes (3). A strong antitumor efficacy of
rIL-12 or of cells engineered to produce IL-12 has been reported in a
great variety of tumors (4-11). Antitumor activity of IL-12 is thought
to be mediated mainly by in vivo induction of IFN--y, given that the
therapeutic efficacy of IL-12 is reduced in mice treated with anti-
IFN-y antibodies (5). The IFN-inducible chemokine IP10 has been
indicated in turn as a further mediator of the antitumor activity of
IL-12 (7).
In this paper, we show that IL-12 gene transduction of a MHC-
negative murine melanoma clone results in a complete loss of malig
nancy, and that IL-12-transduced cells can be used as a therapeutic
vaccine against métastasesproduced by nontransduced MHC-nega
tive parental cells.
MATERIALS AND METHODS
Cells. B78H1 is an amelanotic clone of B16 melanoma (12). B78H1 cells
were cultured in DMEM (Lite Technologies. Inc.. Milan. Italy) supplemented
with 10% fetal bovine serum (Life Technologies. Inc.).
Gene Transduction. IL-12-producing B78H1 cells were obtained by trans
duction with the polycistronic Lp4()Ip'5SN retroviral vector coding for both
p35 and p4() murine IL-12 subunits as described previously (10). Single
G418-resistant colonies were isolated, expanded into cell lines, and subjected
to further analysis. The IL-12 concentration was determined by a two-site
sandwich ELISA using 9A5 mAb to p70 and peroxidase conjugate mAb
5C3-POD to p40, kindly provided by Dr. Luciano Adorini (Roche Milano
Ricerche. Milan. Italy).
Mice. Eight-week-old C57BL/6NCrlBR (referred to as C57BL/6) male
mice and 5-week-old nu/nu male mice on Swiss GDI background were
purchased from Charles River Laboratories (Calco. Italy) and treated according
to European Community guidelines. To obtain NK-depressed animals, we
injected some groups of mice i.v. with 0.4 ml of a 1:25 dilution of anti-asialo
CM, antiserum (WAKO. Dusseldorf. Germany) 24 h before cell injection.
Some mice were treated every 3 days with anti-asialo GM, at the concentration
described above and with 100 /j.g of anti-IFN-y mAb (AN 18 hybridotna;
kindly provided by Dr. G. Carotta. Roche. Basel. Switzerland).
Tumor Growth and Metastasis. Mice were challenged s.c. with 0.2 ml of
a single-cell suspension containing 5 X IO5 (C57BL/6) or IO6 (nude mice)
tumor cells: cell doses were chosen to obtain tumors with a similar incidence
and growth curve in C57BL/6 and in nude mice. The incidence and the growth
of tumors were evaluated twice weekly. Neoplastic masses were measured
with calipers: tumor volume was calculated as 77/6 X [(V (a x b)]}. in which
o and h are two perpendicular major diameters. Experimental métastaseswere
evaluated 28 days after the injection in a lateral tail vein of 5 x IO5 (C57BL/6).
IO6 (nude mice), or 2.5 x IO5 (anti-asialo GM,-treated C57BL/6) tumor cells
suspended in PBS. Lung nodules were contrasted with black India ink; all
metastasis counts were performed on dissected lung lobes under a stereoscopic
microscope.
Therapeutic Vaccination. Therapeutic vaccination started I day after
B78H1 parental cell challenge and consisted of seven injections of 10" viable
or mitomycin C (80 fig/ml at 37°Cfor 45 min: Sigma)-treated cells 3-4 days
apart.
Administration of Exogenous Mouse IL-12. Murine rIL-12. kindly pro
vided by Dr. Maurice Gately (Hoffmann LaRoche. Nutley, NJ). was injected
(0.1 /j.g/day diluted in PBS containing 100 fig/ml murine serum albumin) i.p.
or at the site of s.c. vaccination. Starting 1 day after i.v. injection of tumor
cells, mice received two courses of five daily i.p. injections with a 2-day
1225
Research.
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