[CANCER RESEARCH 37, 1644-1648, June 1977]
SUMMARY
Some of the dynamics of neoplastic transformation in
vitro have been studied with the use of benzo(a)pyrene as
the carcinogen in the C3H/1OT1/2 morphological transfor
mation assay. Experiments that involved the dispersion of
cells into new culture dishes at various times after carcino
gen treatment have shown that no change in the fraction of
potentially transformed cells occurs while cultures grow to
form a confluent monolayer, that little or no change in the
fraction of potentially transformed cells occurs for approxi
mately 3 weeks after confluence is attained, and that this
fraction increases rapidly some 7 weeks after BP treatment.
When confluent benzo(a)pyrene-treated cultures are dis
persed in new culture dishes prior to the onset of growth
toward focus formation , the formation of transformed foci is
suppressed at high cell densities of seeding. This phenome
non is independent of the total number of divisions under
gone by cells after treatment. We suggest that phenotypic
expression of morphological transformation is dependent
on colony interactions in the C3H/1OT1/2 system, which we
do not yet understand, but which are independent of time
posttreatment expressed either in cell generations or abso
lute time.
INTRODUCTION
C3H/1OT1/2 clone 8 is a line of mouse embryo fibroblasts,
isolated by Reznikoff et a!. (12), that is subject to a high
degree of density-dependent inhibition of growth. Carcino
genic stimuli [polycyclic aromatic hydrocarbons (ii), X-rays
(14), smoke condensate (3), etc.] are followed by the ap
pearance of foci of multilayered cniss-cross colonies some 6
to 8 weeks after treatment. Transplantation of cells from
such foci to C3H neonatal mice results in tumor formation
(3, 9, 10, 11). Spontaneous appearance offoci is infrequent
(<10@@).It has been shown, however, that normal untrans
formed BALB/3T3 cells give rise to tumors, when implanted
in host animals preattached to a solid substrate (5, 6, 16).
Similar results have also been obtained with the C3H/1OT1/
2 line (C. Heidelberger, personal communication). This
I This work was supportedin part by GrantsNCI5ORO1and CA 15010-
O2ETfrom the National Cancer Institute, Grant NIEHS 5P01 ES 00597 from
The National Institute of Environmental Health Sciences, and by the Massa
chusetts Institute of Technology Undergraduate Research Opportunities Pro
gram.
Received June 21, 1976; accepted February 25, 1977.
raises questions as to the nature of the C3H line, which will
be discussed later. For the purpose of clarity, however, the
term “transformation,― as used in this paper, will refer to
the morphological alteration in cellular growth patterns,
from monolayered and ordered to multilayered and criss
crossed.
Independent of its potential as an assay for carcinogenic
potential of a test agent, the reproducibility of the system
and its adaptability for defined experimental study make it a
useful tool for the characterization of the process of carci
nogenesis in vitro.
Reznikoff et a!. (11) found that increasing initial cell den
sity above 1000 cells/60-mm dish at the time of carcinogen
treatment suppressed the appearance of transformed foci.
This observation suggested 2 possible explanations. First,
close contact between normal and potentially transformed
colonies, which is maximized at high inoculurn concentra
tions, might exert an inhibitory effect on the production of
transformed foci. Second, cells seeded at high density re
quire fewer divisions to form a confluent monolayer relative
to cells seeded at low density. Thus, the observed inhibition
might reflect a requirement for a particular number of cell
divisions before neoplastic expression would be possible.
Cells treated with carcinogens (8, 9), oncogenic viruses
(2), and X-rays (7, 14) have been reported to require a
minimum number of cell generations before transformation
could be observed. This requirement is now understood as
2 separate requirements, 1 for initial permanent fixation of
the transformed state (usually 1 cell division), without which
the capacity to express transformation is rapidly lost and,
the 2nd, a growth requirement hypothetically needed for
phenotypic expression (8, 9). The 1st cell division may be
needed to fix a potential carcinogenic event. The additional
generations could provide a means by which a key factor no
longer synthesized is reduced to a low concentration
through dilution. For example, expression of phage resist
ance mutations in bacteria may require as many as 14
generations after treatment (13). Expression of the 6-thio
guanine-resistant state in human lymphoblasts also re
quires some 14 generations (15). The experiments de
scnibed in this paper were designed primarily to determine
whether the initial density-dependent inhibition of focus
formation observed by Aeznikoff et a!. was due to the inter
action of potentially transformed colonies with other cob
nies or whether it was the result of a large number of cell
generations required for the expression of carcinogen-in
duced transformation, i.e., a long phenotypic lag.
CANCERRESEARCHVOL. 37
1644
Cell Density Dependence of Focus Formation in the C3H/1OT1/2
Transformation Assay1
Daniel A. Haber, David A. Fox, William S. Dynan, and William G. Thilly
Department of Nutrition and Food Science, Massachusetts Institute of Technology, Cambridge, Massachusetts 02139 fD. A. H. , W. G. T.J; Harvard Medical
School, Boston, Massachusetts 02115 (D. A. F.J;and McArdle Laboratory for Cancer Research, University of Wisconsin, Madison, Wisconsin 53706 fW. S. D.J
on May 5, 2021. © 1977 American Association for Cancer Research. cancerres.aacrjournals.org Downloaded from