ICANCER RESEARCH 56. 4430-4437. October I, IW6|
Correlations between S and G2 Arrest and the Cytotoxicity of Camptothecin in
Human Colon Carcinoma Cells'
Francois Goldwasser,2 Tsunehiro Shimizu, Joany Jackman, Yuko Hoki, Patrick M. O'Connor, Kurt W. Kohn, and
Yves Pommier3
lÂihortiltn~\ of Molecular Pììtinnucolo^y. Division oj Basic Sciences, National Cancer Institute. N1H, Bctlicsilu. Man-land 20ÃOEW1-4255
ABSTRACT
Previous cell line comparisons indicated that neither S-phase fraction
nor topoisonu'rase I {topI) levels are sufficient to predict camptothecin
(CRT) cytotoxicity (F. Goldwasser et al.. Cancer Res., 55: 2116-2121.
1995.). To identify new determinants for CPT activity, two mutant p53
human colon cancer cell lines, SW620 and KM 12, that were previously
reported to have similar topi levels and differential sensitivity to CPT
were studied. No difference in the kinetics of topi-mediated DNA single-
strand hreaks or DNA synthesis inhibition were observed after l h expo
sure to 1 /KMCPT. Pulse-labeling alkaline elution showed deficiency of
damaged replicón elongation in the more sensitive SXV620 cells. Consis
tently, flow cytometry analyses showed that KM12 was arrested in G2,
whereas SW620 cells were irreversibly blocked in S phase. Aphidicolin
protection was minimal in KM 12 and more pronounced in the more
sensitive SYV620 cells. Thus, CPT appears to have two cytotoxic mecha
nisms, one protectable by aphidicolin and present in SVV620 and the other
not protectable by aphidicolin and common to both cell lines. SYV620
exhibited also a greater capacity to break through the I., checkpoint after
DNA damage. Consistently, SVV620 cells failed to down-regulate cyclin
B-cdc2 kinase activity, whereas KM 12 cells down-regulated cyclin B/cdc2
kinase activity within 30 min to 20% of control level after CPT treatment.
Analysis of the 7 human colon carcinoma cell lines of the NCI Anticancer
Drug Screen showed that defects in replicón elongation and (., break
through capability correlate with sensitivity to CPT. Our results suggest
that misrepair of damaged replicons and/or alterations in DNA damage
checkpoints is critical to defining chemosensitivity to CPT-induced topl-
cleavahle complexes and that CPT appears to have two cytotoxic mecha
nisms, one protectable by aphidicolin, and the other not.
INTRODUCTION
CPT4 derivatives exhibit promising activity for the treatment of
solid tumors, such as lung and colon cancers, two high-incidence
diseases with poor prognosis (1—5).CPT and its derivatives are
specific inhibitors of eukaryotic DNA topi (6, 7). a ubiquitous en
zyme with key roles in DNA replication, transcription, and possibly
recombination and repair (for recent review, see Ret. 8). Topi cata
lyzes linking number changes of DNA in steps of one by breaking and
resealing a single phosphodicster bond at a time, and CPTs covalently
trap these top I-DNA intermediates, termed cleavable complexes,
thereby converting topi into a cellular poison (7. 8). Collisions
between advancing replication forks and trapped top I-cleavable com
plexes are considered a critical step for CPT cytotoxicity (9-11)
because they lead to DNA fork breakage and highly toxic DNA DSBs
Received 1/18/96: accepted 7/29/6.
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 in part by a fellowship from the French Ligue Nationale contre le Cancer
and from the NCl/Huropean Organisation for Research and Treatment of Cancer (to
F. G.).
- Present address: Hôpital Paul Brousse. 948IX) Villejuif. France.
3 To whom requests for reprints should be addressed, at Laboratory of Molecular
Pharmacology. Building 37. Room 5C25. National Cancer Institute. NIH. Bethesda. MD
20892-4255. Fax: (301)402-0752.
4 The abbreviations used are: CPT. camptothecin: topi, topoisomcrase I: DSB. double-
strand break; SSB. single-strand break: NCI. National Cancer Institute; GI„„. 90'/r cell
growth inhibition.
(12. 13). This DNA-damaging mechanism, at least in some cell types,
accounts for the maximum toxicity of CPT derivatives during S phase
despite nearly constant topi protein levels throughout the cell cycle
(II. 14).
CPT cytotoxicity also depends on the potential of a cell to undergo
apoptosis or respond adequately to DNA damage (11). Replication-
dependent DNA strand breaks induce wild-type p53 protein (15),
which may lead to arrest at the G,-S transition and prevention of "cell
commitment and progression through S phase" ( 16). Arrest in G-, is a
common cellular response to DNA damage ( 17-19); it is mediated by
a negative feedback control; it is also referred to as a checkpoint
response to DNA damage (20-22). Down-regulation of cyclin B/cdc2
activity after CPT exposure arrests cells in G, and presumably allows
DNA repair before commitment into M phase (23). To identify
determinants for CPT activity, we previously used the colon cancer
cell lines of the NCI Anticancer Drug Screen as a "miniclinic" to
identify cell lines that exhibit intrinsic sensitivity and resistance to
CPTs ( 14). In contrast to hematopoietic cells (24). colon cancer cells
do not undergo rapid apoptosis.5 Our results showed that the seven
colon carcinoma cell lines of the NCI cell screen exhibit a range of
CPT sensitivity that is better correlated with the frequency of cleav
able complexes than with topi levels ( 14). Two of these lines. SW620
and KM 11. both with mutant p53 genes'1 responded very differently to
CPT despite similar levels of topi expression and top I-cleavable
complexes (14).
The aim of the present study was to identify determinants for CPT
activity located downstream from topi-cleavable complexes, using
these two colon cell lines, SW620 and KM 12. and to correlate these
determinants and cytotoxicity using the overall panel of colon cell
lines from the NCI Anticancer Drug Screen. Our results suggest that
CPT cytotoxicity is related to two types of defects: (a) during S phase,
accumulation of damaged replicons and failure to complete elongation
of replicating DNA; and (/>) during G-,. deficiency to down-regulate
cyclin B/cdc2 kinase and reduced G2 arrest. We also report an S
phase-independent CPT-induced cytotoxicity.
MATERIALS AND METHODS
Cell Culture. SW620. KMI2. HCT1I6. HCTI5. HCC2998. HT29. and
coli>2()5 correspond to the seven colon cancer cell lines used in the NCI
Anticancer Drug Screen (25). They were obtained from Dr. Dominic Scudieri)
(NCI, Frederick. MD). Cells were grown at 37°Cin 95% air/5% CO, in RPM1
1640 (Lite Technologies. Inc.. Grand Island. NY) containing 5r/f fetal bovine
serum (Inlergen. Purchase, NY). 2 HIMglutamine (AB1, Columbia. MD).
Drugs and Chemicals. CPT was a kind gift from Drs. Monroe E. Wall and
Mansukh C. Wani (Research Triangle Institute. Research Triangle Park. NC).
Nocoda/.ole was purchased from Sigma Chemical Co. (St. Louis, MO). Stock
solutions of CPT and nocodazole were prepared in DMSO at 10 HIMand 10
mg/ml, respectively, and further diluted in deionized water immediately prior
to each experiment. [y^PI ATP (4500 mCi/mmol) was purchased from NEN
Research Products (DuPont NEN. Boston. MA).
s F. Goldwasser and Y. Pommier, unpublished data.
" P. M. O'Connor. I. Jackman. I. Bae. T. G. Myers. S. Fan. D A. Scudieri). A. Monks.
E. A. Sausville, J. N. Weinstein. S. Friend. A. J. Fornace. Jr.. and K. W. Kohn. manuscript
in preparation.
4430
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