(CANCER RESEARCH 46, 2863-2865, June l986J
studies have used the two-stage tumorigenesis protocol, concen
trating on events during initiation and promotion. Since carci
noma development is relatively rare (approximately 10% of
papillomas progress to carcinomas) and carcinoma is a late
event in the two-stage protocol, little has been reported on the
modification of malignant progression. The observation that
carcinomas occur relatively frequently in models in which car
cinogens are applied repetitively has led to the proposal that
progression is closely related to initiation (6). Several investi
gators have treated mouse skin papillomas with tumor initiators
and observed increased carcinoma formation compared to con
trol mice treated with the potent tumor promoter 12-O-tetra
decanoylphorbol-13-acetate (7, 8). These researcherssuggested
that progression to malignancy was the result ofgenetic changes
causedby mutagenic agents.
Benzoyl peroxide is a free radical generating agent, used as
an industrial polymerizing agent, as a curing agent for flour
and cheeseand asan additive in cosmetics and pharmaceuticals,
including those used in the treatment of acne (9, 10). It is
moderately active as a tumor promoter but is inactive as a
complete carcinogen or tumor initiator (1 1, 12). Similar to
other tumor promoters, benzoyl peroxide induces an increase
in dark basal keratinocytes, epidermal hyperplasia, terminal
differentiation, and ODC levels, and it inhibits intercellular
communication in both mouse (12, 13) and human cells (14,
15). As it is true for other agents, these promotional effects
may be related to benzoyl peroxide's free-radical generating
ability (12). In previous work (16) tumor promotion with ben
zoyl peroxide (20 mg twice weekly) was found to cause an
unexpectedly high rate of carcinomas relative to papillomas.
We hypothesized that in addition to its promotional activity
benzoyl peroxide may be highly active in malignant progression,
the third stage of carcinogenesis. This report shows that some
tumor promoters have the ability to alter malignant progres
sion.
MATERIALS AND METHODS
Female SENCAR mice were receivedat 5—7wk of age (Harlan
Laboratories, Indianapolis, IN). Skin tumors were chemically induced
using a standard two-stage initiation-promotion protocol. Initiation
was accomplished by a single topical application of 10 nmol DMBA4
(Aldrich, Milwaukee, WI) on the shaveddorsal skin. Biweekly promo
tion with 1 ;zgof 12-O-tetradecanoylphorbol-l3-acetate (L. C. Services
Corp., Worburn, MA) was begun 2 wk after initiation. All solutions
were applied in 0.2 ml of acetone.
Beginning with wk 21, one group of papilloma-bearing mice (21
animals) continued to receive twice-weekly applications ofTPA (1 gig),
while a second group (20 animals) was begun on twice-weeklytreat
ments of 20 mg of benzoyl peroxide. Treatments were continued until
theterminationof theexperimentat wk 40.
Papilloma yield between groups was expressed as the number of
papillomaspermouse±SEandcomparedbyStudent'st test.Carci
nogenesis was followed by recording the cumulative number of mice
whichdevelopedcarcinomasandthecumulativenumberofcarcinomas
4 The abbreviations used are: DMBA, 7,12-dimethylbenzanthri
O-tetradecanoylphorbol-13-acetate; GGT,-y-glutamyltransferase.
ABSTRACF
Chemical carcinogenesis in mouse skin can be divided into the proc
esses ofinitiation, promotion, and progression. The free-radical generator
benzoyl peroxide is moderately active during the promotion stage. Re
petitive treatment ofmouse benign skin tumors (papillomas) with benzoyl
peroxide (20 mg, twice weekly) increased the number of cumulative
carcinomas per group by 325% and the number of keratoacanthomas by
44% compared to tumor-bearing Sencar mice treated with the promoter
12-O-tetradecunoylphorbol-13-acetate. The lack of increase in the num
her ofcumulative papillomas per group due to benzoylperoxidetreatment
suggests that benzoyl peroxide enhanced the progression of preexisting
papillomas. The ability of benzoyl peroxide to enhance the progression
of benign tumors to cancer should be considered when determining the
human risk from exposure to this widely used chemical agent in addition,
biological assays specifically testing malignant progression may be es
sential and beneficial for determining an agent's carcinogenic risk.
INTRODUCFION
Carcinogenesis in several experimental systems can be di
vided into processes of initiation, promotion, progression, and
metastasis. The separation between initiation and promotion
has been clearly demonstrated in mouse skin (1) in which a
single subthreshold dose of a carcinogen (initiation) followed
by repetitive applications of a noncarcinogenic, nonmutagenic
agent (promotion) produces first papillomas and then squamous
cell carcinomas (two-stage tumorigenesis). Tumors that arise
from repetitive applications of a carcinogen (complete carci
nogenesis) are theoretically the result of both the initiating and
promoting abilities of most carcinogens.
Evidence indicates that initiation involves the induction of
permanent genetic alterations and is essentially irreversible.
Most, but not all, initiators are classified as mutagens in stand
ard in vitro mutagenesis assays (2). Promotion seems to involve
clonal expansion of initiated cells leading to the expression of
benign tumors. In mouse skin, promotion can be subdivided
into two stages: an early stage that correlates with dark basal
keratinocyte induction and a late stage that correlates with
epidermal hyperplasia and ornithine decarboxylase induction
(3, 4). The biological significance of many of the biochemical
and molecular events during promotion is not clear. Progres
sion, the process by which a benign tumor develops malignant
characteristics is proposed to involve a number of genetic
alterations. In addition to their invasive behavior, mouse skin
squamous cell carcinomas appear morphologically and geneti
cally more deviant than either normal or benign tissue (5).
Squamous cell carcinomas of mouse skin rarely metastasize to
other organs, but their metastasis can be increased by applica
tions of alkylating agents.3
During the last two decadesmost mouse skin carcinogenesis
Received 11/4/85; revised 2/10/86; accepted 2/25/86.
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
accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
I These studies were supported by NIH grants CA-34890, CA-34962, CA
34521,andCA 38863.
2 To whom requests for reprints should be addressed.
3 1. J. Slaga, unpublished results.
TPA, 12-
2863
Enhanced Malignant Progression of Mouse Skin Tumors by the Free@Radical
Generator Benzoyl Peroxide'
John F. O'Connell,2 Andres J. P. Klein-Szanto, Donna M. DiGiovanni, JoAnn W. Fries, and Thomas J. Slaga
The University ofTexas System Cancer Center, SciencePark-ResearchDivision, Smithvile, Texas 78957
Research.
on January 22, 2022. © 1986 American Association for Cancer cancerres.aacrjournals.org Downloaded from