(CANCER RESEARCH 52, 4922-4928, September 15. 1992|
Elevation of Thymidylate Synthase following 5-Fluorouracil Treatment Is
Prevented by the Addition of Leucovorin in Murine Colon Tumors1
Clasina L. Van der Wilt, Herbert M. Pinedo, Kees Smid, and Godefridus J. Peters2
Department of Oncology, Free University Hospital, 1007 MB Amsterdam, The Netherlands
ABSTRACT
The inhibition of thymidylate synthase (TS) by the 5-fluorouraciI
(5FU) metabolite 5-fluorodeoxyuridine monophosphate can be biochem
ically modulated by leucovorin (LV). LV administration increases the
level of reduced folates in tissues, which promotes the inhibition of TS.
We have studied the antitumor effect, free 5-fluorodeoxyuridine mono-
phosphate levels, and inhibition of TS in two murine colon tumors at
several time points after weekly 5FU or LV and 5FU administration.
The antitumor effect of 5FU alone could be potentiated by LV in both
tumors. 5-Fluorodeoxyuridine monophosphate levels (212 and 46
pmol/g wet wt. after 2 h for Colon 26 and Colon 38, respectively) were
sufficient to mediate TS inhibition, but the levels were not related to
antitumor activity. Untreated controls of the 5M -sensitive tumor Colon
38 had 3 times lower TS levels than did those of the more resistant
tumor Colon 26. One course of treatment resulted in a comparable
extent and retention of TS inhibition for 511 and LV/5FU therapy in
both tumors. After 1 week there was complete recovery of TS inhibition,
but the TS levels in tumors from 5FU-treated mice tended to be higher
than the controls, which was more pronounced after three courses of
therapy. A 4-fold increase of TS levels was seen in Colon 26 after SFU
therapy. The elevation of TS in this tumor affected the extent of TS
inhibition. Tumors treated with 5FU and LV also showed an increase of
TS, but to a lesser extent, while the absolute effect on TS inhibition
remained the same. This might be related to the potentiating effect of
LV on 5FU antitumor activity in vivo in these tumors.
INTRODUCTION
The enzyme TS3 is considered to be a major target of 5FU,
which is commonly used in the treatment of advanced colorec-
tal cancer. TS catalyzes the reductive methylation of dUMP,
which is the final reaction in the de novo synthesis of dTMP.
Inhibition of TS leads to depletion of dTMP and, thereby, to
cessation of DNA biosynthesis. The inhibition of TS is medi
ated by the formation of a ternary complex consisting of TS,
FdUMP, and the cofactor CH2-THF (1). The availability of
CH2-THF, especially in its polyglutamated form, determines
the stability of the ternary complex (2-4), and limiting amounts
might affect the action of 5FU on TS. This is the rationale for
the combined treatment with 5FU and LV, a precursor of CH2-
THF.
In vitro (5-7) and in animal models (8-10), LV potentiated
the effect of 5FU. In patients, it has also been demonstrated
that LV can enhance the response rate of single-agent 5FU, in
advanced colorectal cancer, from <20% to 30-40% (11). An
Received 12/20/91; accepted 7/3/92.
The costs of publication of this article were defrayed in part by the payment of
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ance with 18 U.S.C. Section 1734 solely to indicate this fact.
1This study was supported by The Netherlands Cancer Foundation (Koningin
Wilhelmina Fonds Grant IKA-VU 88-20) and Lederle Cyanamid, The Nether
lands.
2 To whom requests for reprints should be addressed, at Department of Oncol
ogy, Free University Hospital, P.O. Box 7057, 1007 MB Amsterdam, The Neth
erlands.
3 The abbreviations used are: TS, thymidylate synthase; CH2-THF, 5,10-meth-
ylenetetrahydrofolate; 5FU. 5-fluorouracil; LV, leucovorin; FdUMP, 5-fluorode
oxyuridine monophosphate.
effective schedule now used clinically (12) is comparable to the
most effective schedule in mice, consisting of pretreatment with
LV followed by simultaneous administration of LV and 5FU
(10). It is generally believed that this potentiation is due to
enhancement of TS inhibition and that the degree and duration
of TS inhibition induced by 5FU treatment might influence the
success of treatment.
Measurements of TS before and after exposure to 5FU have
been performed in vitro (13, 14). TS levels and TS inhibition
after treatment have also been measured in tumor tissue of
animal models (15-17) and patients (18-22). Most studies con
centrated on the effect of a single dose of 5FU on TS levels,
which was evaluated for a maximum of 72 h (16), while enzyme
assays were usually limited to the ligand binding assay with
FdUMP.
We studied the long term effects of 5FU and LV/5FU treat
ment in two murine colon tumors, Colon 26 and Colon 38.
These tumors have been characterized extensively with respect
to 5FU metabolism and response to 5FU-containing regimens
(10, 23-25). Colon 38 is more sensitive to 5FU than is Colon
26. Potentiation of the antitumor effect of 5FU by LV appeared
to be schedule dependent, and the combination was more effec
tive when LV was administered before and together with 5FU
(10). Recently, a purified stereoisomer of LV, /-LV, has become
available. This is thought to be the biologically active form of
LV and might, therefore, have a better effect than the racemic
mixture of d- and /-LV. The clinical effects of/-LV are currently
being studied (26, 27).
TS inhibition after 5FU treatment has been studied for short
periods, but little is known about long term effects. Several
mechanisms of resistance related to TS (19) might, however, be
present in the tumor and could be activated or develop during
therapy. This study concentrates on the effects of 5FU treat
ment on TS after more than one treatment course, since pa
tients usually receive more than one course of 5FU administra
tion. The present results demonstrate that prolonged treatment
with 5FU can induce an increase in TS levels, which can, how
ever, be prevented by concomitant LV therapy.
MATERIALS AND METHODS
Chemicals. 5FU was obtained from Hoffman LaRoche (Basel, Swit
zerland). Leucovorin and purified /-leucovorin were kindly provided
by Lederle Cyanamid (Etten-Leur, The Netherlands). [6-3H]-FdUMP
(specific activity, 20 Ci/mmol) was purchased from Moravek Bio-
chemicals Inc. (Brea, CA) and [5-3H]-dUMP (specific activity, 10.9
Ci/mmol) from Amersham International (Buckinghamshire, England).
Lactobacillus casei thymidylate synthase (purified enzyme) was ob
tained from BIOPURE (Boston, MA). ¿/-Tetrahydrofolate (Sigma
Chemical Co., St Louis, MO) was converted into CH2-THF by addition
of formaldehyde (15, 28). All other chemicals were of analytical grade
and were commercially available.
In Vivo Studies. Maintainance of the tumors in mice and treatment
with 5FU and LV have been described in detail previously (9, 10). The
most effective schedule for the combined treatment with LV and 5FU
was found to be 50 mg/kg LV, followed, after 1 h, by 50 mg/kg LV and
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