[CANCER RESEARCH 46, 1754-1758, April 1986]
Noninvasive Spectroscopic Analysis of Fluoropyrimidine Metabolism in Cultured
Tumor Cells1
Max Keniry, Chris Benz,2 Richard H. Shafer, and Thomas L. James
Departments of Pharmaceutical Chemistry [M. K., R. H. S., T. L. JJ and Radiology ¡T.L. J.J and Cancer Research Institute [C. B.], University of California,
San Francisco, California 94143
ABSTRACT
Nuclear magnetic resonance spectroscopy is a technique that may be
used noninvasively to follow the intracellular metabolism of fluorinated
antimetabolites such as 5-fluorouracil (FUra) and S-fluorouridine. Intra
cellular "F spectral peaks are assigned by comparison with the pH-
dependent chemical shifts measured for eight commercially available
fluoropyrimidine metabolites as well as by comparison with the literature
recorded values of five known catabolites of FUra. Five murine and
human tumor cell lines (N,S„ Sarcoma 180, LI210, HI,-60, and Mia-
PaCa) were exposed in vitro for 24 h to cytostatic doses of FUra or 5-
fluorouridine. Treated cells were harvested and analyzed immediately or
following a subsequent incubation under either nutrient-rich or nutrient-
poor conditions. A major narrow component peak at 4.6-4.9 ppm was
observed in all cell samples analyzed immediately after treatment. This
peak was identified as intracellular FUra nucleotides, and its T, value
was approximately 800 ms. No fluoropyrimidine catabolites were detect
able in any of the treated cell lines. Free FUra could be measured in cells
only after subsequent incubation under nutrient-poor conditions, and this
was associated with a decline in the prominent FUra nucleotide peak. In
treated cells chased with drug-free media containing 1 pM thymidine,
spectra revealed a broad component signal underlying and downfield from
the narrow nucleotide-containing peak. By biochemically fractionating
treated cells into an acid-soluble fraction and phenol-purified cytoplasmic
and nuclear RNA extracts, we were able to completely separate the
nucleotide peak from the broad component signal resulting from FUra
incorporation into RNA. Thymidine produced a marked enhancement of
this "F signal into both cytoplasmic and nuclear RNA without affecting
the nucleotide signal from the acid-soluble fraction. The present ability
of nuclear magnetic resonance to monitor the metabolic channeling of
fluoropyrimidines in intact tumor cells suggests that future spectroscopic
imaging of patients treated with fluorinated antimetabolites may provide
clinically important information about tumor biochemistry and drug
sensitivity.
INTRODUCTION
Fluorinated pyrimidines are antimetabolites frequently used
in the treatment of breast and gastrointestinal cancers (1); they
are also occasionally used in the treatment of refractory lym-
phomas and leukemias (2). The metabolism of these drugs has
been extensively studied in murine and human tumor cell lines,
and it now is well established that the rate limiting intracellular
uptake of FUra,3 FUrd, or FdUrd is determined by three
individual phosphorylating enzymes involved in pyrimidine
synthesis (3-5). Once phosphorylated in the cell, the three
fluoropyrimidines may interconvert or become anabolized into
the primary cytotoxic nucleotides, FUTP and FdUMP. The
former ribonucleotide is incorporated into RNA, disrupting
Received 8/14/85; revised 12/30/85; accepted 1/2/86.
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.
1Supported in part by grants CA-36769, CA-36773, and CA-27343 from the
National Cancer Institute and CH-235B and NP-437 from the American Cancer
Society.
2To whom requests for reprints should be addressed.
3 The abbreviations used are: FUra, 5-fluorouracil; FUrd, 5-fluorouridine;
FdUrd, 5-fluorodeoxyuridine; FdUMP, 5-fluorodeoxyuridine monophosphate;
FUDP, 5-fluorouridine diphosphate; FUTP, 5-fluorouridine triphosphate; NMR,
nuclear magnetic resonance; FUMP, fluorouridine monophosphate.
RNA processing and protein synthesis; the latter deoxynucleo-
tide is a suicide inhibitor of thymidylate synthetase, an enzyme
essential for DNA synthesis. The relative importance of these
two anabolhes in the clinical antitumor effects of FUra remains
a matter of dispute (6-8).
Virtually all available data on the intracellular metabolism of
fluoropyrimidines derive from the biochemical analysis of bro
ken (lysed) cell preparations, usually following addition of some
radiolabeled precursor. NMR is an alternative technique that
may be used noninvasively to follow the metabolism of natural
"F-containing compounds. We and others have shown that it
is possible to monitor fluoropyrimidine metabolism within bac
terial cells (9) or from the biological fluids of drug-treated
animals (10, 11). The present study extends these results by
utilizing "F-NMR to monitor in vitro the metabolism of FUra
and FUrd in a variety of murine and human tumor cell lines.
MATERIALS AND METHODS
Drugs, Cells, Cytotoxicity, and Biochemical Assays. The commer
cially obtained fluoropyrimidines used in this study were FUra, FUrd,
FdUMP, 5-fluorouridylate, 5-fluorocytosine (Sigma, St. Louis, MO),
5-fluorocytidine (Hoffman-LaRoche, courtesy of A. F. Cook), 5-fluo-
roorotate (Pharmacia, NJ), and FUTP (Sierra Bioresearch, Tucson,
AZ).
Tumor and leukemia cell lines were maintained as stock cultures in
5% CO2 incubators at 37°Cusing RPMI 1640 medium (Gibco Labo
ratories, NY) supplemented with 5-10% fetal calf serum (HyClone
Laboratories, UT). The NtSi (Novikoff). Sarcoma 180, and L1210 are
well-characterized, rodent-derived hepatoma, sarcoma, and leukemia
cell lines, respectively, which grow as cell suspensions in vitro. The
MiaPaCa cells are a long-established monolayer cell line derived from
an undifferentiated human pancreatic carcinoma, and the HI. 60 cells
are a suspension culture of a widely available human promyelocytic
leukemia cell line. For drug treatment prior to NMR analysis, or for
assessment of drug Cytotoxicity, exponentially growing cells (in 75-enr
sterile plastic flasks: Costar, Cambridge, MA) were exposed for 24 h to
either FUra or FUrd at the indicated concentrations. All the cell lines
studied were growth inhibited by 24-h exposure to either 100 MMFUra
or 2.5 tiM FUrd. Complete growth curves were obtained for treated
NiSi and Sarcoma 180 cells, as shown in Fig. 1. In most experiments,
thymidine (1 pM) was added to the culture media 6 h following the
addition of fluoropyrimidine to circumvent FdUMP inhibition of DNA
synthesis and maximize incorporation of FUTP into RNA. After 24 h,
the cells were harvested, rinsed free of drug, and then resuspended in
fresh, thymidine-supplemented media. To measure the growth-inhibit
ing effect of drug exposure, treated and untreated cells were then
counted on a Coulter Counter (Coulter Electronics, Inc., Hialeah, FL)
at frequent intervals for an additional 96 h, and these cell counts were
plotted on log-linear graphs. For NMR analysis, the freshly harvested
cells were chilled to 4*C and placed in a glass 10-mm diameter NMR
tube for immediate spectroscopy. Biochemical fractionation was per
formed on harvested Sarcoma 180 cells. Pellets of approximately 5x
10* cells were either treated with 0.5 M cold perchloric acid resulting
in an acid-soluble extract containing catabolites, bases, nucleosides,
and nucleotides (12) or alternatively, the chilled cells were gently lysed
and separated into nuclear and cytoplasmic fractions for RNA purifi
cation.
Isolation and Purification of Cytoplasmic and Nuclear RNA. Sarcoma
1754
Research.
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