[CANCER RESEARCH 47, 3212-3219, June 15, 1987]
Activation of the K-ras Protooncogene in Lung Tumors from Rats and Mice
Chronically Exposed to Tetranitromethane
S. Jill Stowers,1 Paul L. Glover, Steven H. Reynolds, Lawrence R. Boone, Robert R. Maronpot, and
Marshall W. Anderson
Laboratory of Biochemical Risk Analysis, Biometry and Risk Assessment Program fS. J. S., S. H. R., M. W. A.], Cellular and Genetic Toxicology Branch [P. L. G.,
L. R. B.I. and Chemical Pathology Branch [R. R. M.], Toxicology Research and Testing Program, National Institute of Environmental Health Sciences, Research
Triangle Park, North Carolina 27709; and Toxicology Program, North Carolina Slate University, Raleigh, North Carolina 27650 fS. J. S.J
ABSTRACT
Dominant transforming genes were detected in lung tumors from
Fischer 344 rats and C57BL/6 x C3H I , mice chronically exposed by
inhalation to tetranitromethane, a highly volatile compound used in
several industrial processes. The rat lung neoplasms were classified as
adenocarcinomas, squamous cell carcinomas (epidermoid carcinomas), or
adenosquamous carcinomas. The mouse lung tumors were classified as
papillary adenocarcinomas or adenomas. In both species, the tumors were
morphologically similar to lung tumors in humans. The transfection assay
using NIH/3T3 mouse fibroblasts detected transforming genes in 74%
(14 of 19) of the rat lung tumors and in 100% (4 of 4) of the mouse lung
tumors. Southern blot analysis indicated that transforming gene was an
activated K-ras protooncogene in both species. The first exon of the k-
ras gene in normal DNA and in DNA from two cell lines transformed by
tumor DNA was compared by cloning and sequencing the gene. Experi
ments showed that there was a (.(—»Al transition in the second base of
the 12th codon of the k-ru.v oncogene in the two transfectant DNAs.
Oligonucleotide hybridization indicated that all of the rat and mouse
transfectants had this activating lesion. Additional tumor DNA was then
tested for the presence of a mutated alÃ-elewith the GC—»AT transition.
All of the rat tumors tested and all of the mouse tumors tested had this
mutation present. Hybridization using the normal oligonucleotide se
quence around the 12th codon indicated that the normal alÃ-elewas also
present in the majority of the tumors, suggesting that the loss of normal
alÃ-ele is not necessary for the development of neoplasia. One rat lung
tumor had no normal alÃ-ele present, possibly suggesting that this tumor
could have been in a more advanced stage than the other tumors. This is
the first study to detect activated protooncogenes in rodent tumors induced
under conditions which mimic human exposure to a chemical in the
workplace. Tetranitromethane may exert its carcinogenic action by both
activation of the K-ra\ oncogene and stimulation of cell proliferation by
its irritant properties.
INTRODUCTION
Recent studies suggest that the activation of protooncogenes
by genetic alterations may play a role in leading a cell to
neoplastic development. These genetic alterations include gross
chromosomal rearrangements, amplification of genes, and
point mutations. Oncogenes that have been shown to acquire
transforming activity by point mutation in their coding se
quence include members of the ras oncogene family, the H-ras,
K-ras, and the N-ras (1-13) and the neu oncogene (14). The
activation of the raÃ-family of genes usually occurs via a point
mutation at the 12th, 13th, or 61st codons in human tumors
and tumor cell lines (1-13). Studiesina variety of animal model
systems have shown that specific activation of a protooncogene
by point mutation can be caused by chemical or physical insult
(1-8).
Animal model systems for carcinogenesis have provided a
good means to study protooncogene activation in tumor devel-
Received 11/24/86; revised 3/23/87; accepted 3/25/87.
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1To whom requests for reprints should be addressed.
opment. The H-ras protooncogene has reproducibly been found
activated in rat mammary carcinomas induced by a single
injection of yV-methyl-./V-nitrosourea given during sexual devel
opment (3). The H-ras protooncogene has also been found
activated in mouse skin papillomas and carcinomas induced by
DMBA2 followed by phorbol ester (12-0-tetradecanoylphorbol-
13-acetate) promotion (1, 2, 7). In both models, the H-ras
protooncogene was found to be activated in 90-100% of all of
the tumors examined. Other studies have found K-ras and N-
ras activation in X-ray- or JV-methyl-7V-nitrosourea-induced
mouse thymomas and in rat mesenchymal kidney tumors in
duced by treatment with methyl(methoxy-methyi)nitrosamine
(4, 5, 15). One conclusion from these studies is that exposure
to carcinogens either by relatively high single or multiple doses
causes changes in the DNA resulting in activation of oncogenes.
However, no studies have examined protooncogene activation
in tumors that develop after long term, chronic exposure to
chemicals. The identification of chemicals as potential human
carcinogens is often made on the basis of long term rodent
bioassays which are designed to consider route of human ex
posure and concentrations similar to those present in the envi
ronment, workplace or home.
In a recent bioassay conducted by the National Toxicology
Program, chronic exposure to the industrial chemical TNM
induced a high incidence of primary lung tumors in Fischer 344
rats and C57BL/6 x C3H F, (hereafter called B6C3F,) mice.3
TNM is a highly volatile compound used as a reagent in
industrial nitrosating processes, as an oxidant in rocket fuel,
and as an explosive when mixed with toluene (tetranitro-
toluene). Because of its irritant properties, TNM has also been
proposed as a war gas. The threshold limit for occupational
exposure to TNM based on its irritant properties has been set
at 1 ppm. In the bioassay, groups of 50 male and 50 female
Fischer 344 rats or B6C3F, mice were exposed to TNM by
inhalation for 6 h a day, 5 days a week for 2 years. The rats
were exposed to 0, 2, and 5 ppm of TNM while the mice were
exposed to 0, 0.5, and 1 ppm. Based on histomorphological
examination, the TNM-induced primary lung tumors were ad
enomas, adenocarcinomas, squamous cell carcinomas, and ad
enosquamous carcinomas in rats and papillary adenomas and
adenocarcinomas in mice. These tumors were morphologically
similar to primary lung tumors in humans. The purpose of this
study was to identify and characterize any activated oncogenes
that might be present in lung tumors from rats and mice after
chronic exposure to TNM.
MATERIALS AND METHODS
Lung Tumor Generation. Two-year toxicity and carcinogenicity stud
ies of TNM were performed under National Toxicology Program
2The abbreviations used are: DMBA, 7,12-dimethylbenzanthracene; TNM,
tetranitromethane.
3The National Toxicology Program has not yet completed its evaluation of
the data collected during the studies with TNM. Therefore, the apparent associ
ation of TNM exposure with lung tumors in rats or mice should be considered
preliminary, pending approval of the National Toxicology Program Technical
Report on TNM by the National Toxicology Program Board of Scientific Coun
selors Technical Reports Review Subcommittee.
3212
Research.
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