[CANCER RESEARCH 49, 5907-5912, November 1, 1989]
Independent Mechanisms for Tumor Promoters Phénobarbitaland
12-0-Tetradecanoylphorbol-13-acetate in Reduction of Epidermal
Growth Factor Binding by Rat Hepatocytes1
Sharon A. Meyer, T. Avery Gibbs, and Randy L. Jirtle2
Division of Radiation Oncology, Department of Radiology, Duke University Medical Center, Durham, North Carolina 27710
ABSTRACT
Primary cultures of hepatocytes derived from adult Fischer 344 rats
were used to test for effects of the liver tumor promoter phénobarbital
on several components of the epidermal growth factor (EGF) receptor
signal transduction pathway. Phénobarbitalhad no effect on the binding
of 12SI-labeledEGF by its hepatocyte receptor at 4°Cor on EGF-induced
receptor down-regulation. However, pretreatment of hepatocytes with
phénobarbital (3 mivi)at 37°C caused inhibition of subsequent I25l-labeled
EGF binding. This response temporally resembled that of hepatocytes to
12-O-tetradecanoylphorbol-13-acetate (TPA) in that maximal inhibition
occurred after l h of pretreatment but was reversed after longer pretreat-
ment times. The inhibitory effects of phénobarbitaland TPA on EGF
binding were additive, suggesting that distinct mechanisms mediated the
responses to these two tumor promoters. In addition, treatment with
TPA, but not phénobarbital,caused a redistribution of the activity of
Ca2*/phospholipid-dependent protein kinase C. In untreated and pheno-
barbital-treated hepatocytes, 20% of protein kinase C activity was iso
lated with a membranous fraction, while 75% of the activity was mem
brane associated in TPA-treated hepatocytes. These results demonstrate
that phénobarbital, like TPA and other tumor promoters, can modulate
the EGF receptor system but suggest that it does so without directly
competing with EGF for binding to its receptor or by activating protein
kinase C.
INTRODUCTION
Formation of putative preneoplastic lesions during the pro
motional phase of multistep carcinogenesis involves the expres
sion of a growth advantage by initiated cells relative to normal
cells (1). Thus, one focus of tumor promotion research has been
directed towards describing the effects of tumor promoters on
mitogenic responses of initiated and normal cells to growth
factors and on the signal transduction pathways mediating their
effects. EGF3 is a mitogen for several types of cells (2) and the
pathway mediating EGF-induced mitogenesis has been shown
to be a target of tumor promoter action. The most well-docu
mented effects are those of the phorbol ester class of skin tumor
promoters, e.g., TPA, on the EGF receptor. A rapid, transient
decrease in EGF binding of several types of cultured cells is
generally observed (3-5) and results from phosphorylation of
the EGF receptor as a consequence of TPA activation of Ca2"1"/
phospholipid-dependent protein kinase C (6-8). A similar re
sponse to other structurally distinct classes of skin tumor pro
moters has also been described (9-11), although one of these,
palytoxin, mediates its effects on the EGF receptor independent
of protein kinase C (11).
Experimental carcinogenesis of rodent liver, like that of skin,
is a multistep process, and in this system phénobarbitalis a
Received 2/20/89; revised 7/27/89; accepted 8/4/89.
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.
' This work was supported by USPHS Grants CA25951 and CA40172.
2To whom requests for reprints should be addressed, at Department of
Radiology, Box 3433, Duke University Medical Center, Durham, NC 27710.
'The abbreviations used are: EGF. epidermal growth factor; TPA, 12-O-
tetradecanoylphorbol-13-acetate; MEM, minimal essential medium: EGTA, eth-
ylene glycol bis(f)-aminoethyl ether)-/V,/V,yv',JV'-tetraacetic acid.
strong tumor promoter (1, 12, 13). EGF is a potent mitogen
for rat hepatocytes (14) and phénobarbital, added directly to
the medium of cultured rat hepatocytes, biphasically alters
EGF-dependent mitogenesis. Stimulation has been observed at
phénobarbitalconcentrations of 1 mM or less, while inhibition
is observed at higher concentrations (15, 16). These observa
tions suggest that phénobarbitalmodulates some component(s)
of the EGF-induced mitogenic pathway in hepatocytes. By
analogy to the effect of skin tumor promoters, the hepatocyte
EGF receptor may therefore be a potential site of phénobarbital
action. Also, a recent report (17) describing phénobarbital
inhibition of protein kinase C activity suggests that phénobar
bital could possibly affect EGF binding by modifying protein
kinase C-dependent receptor phosphorylation.
The purpose of the studies described here was to test whether
phénobarbitaladded to the medium of cultured rat hepatocytes
affected early events involved in EGF interaction with its recep
tor. We describe results demonstrating that phénobarbitaldid
not directly compete with EGF for binding to its receptor or
alter EGF-induced receptor down-regulation but that pretreat
ment (37°C)with phénobarbital did cause a rapid, transient
decrease in EGF binding at the hepatocyte surface. Although
this response was temporally similar to that induced by TPA,
further evidence is presented suggesting that the reduction of
hepatocyte EGF receptors by phénobarbital occurred in the
absence of protein kinase C activation.
MATERIALS AND METHODS
Materials. Female Fischer 344 rats (—175g)were from Charles River
Laboratories (Wilmington, MA). MEM (Earle's salts) plus nonessential
amino acids was from GIBCO (Grand Island, NY). Collagenase type I
and receptor grade EGF were purchased from Worthington Biochem-
icals (Freehold, NJ) and Collaborative Research (Bedford, MA), re
spectively. Amersham (Arlington Heights, IL) was the source of 125I-
labeled EGF and [-y-"P]ATP was from NEN-Dupont (Boston, MA).
Sodium phénobarbitalwas from Mallinckrodt (Paris, KY). Leupeptin
was purchased from United States Biochemical Corp. (Cleveland, OH).
Sigma (St. Louis, MO) was the source of bovine serum albumin, bovine
pancreas insulin, histone (type III-S), soybean trypsin inhibitor (type I-
S), phosphatidylserine, TPA, and ff-phorbol.
Hepatocyte Isolation and Culture. Hepatocytes were isolated by in
situ collagenase perfusion, plated (10s cells/cm2) on collagenase-coated
tissue culture plastic, and cultured (37°C, 5% CO2) as described previ
ously (18). Culture medium was MEM with nonessential amino acids
supplemented with pyruvate (1 mM), aspartate (0.2 mM), serine (0.2
mM), insulin (0.1 //M), and gentamicin (50 ¿ig/ml).Fresh medium or
medium containing phénobarbital,TPA, 0-phorbol, or dimethyl sulf-
oxide was applied by replacing the plating medium after allowing 2 h
for hepatocyte attachment, and drug treatments were continued at 37°C
for the times specified. Stock solutions of sodium phénobarbital(10
mM), made on the day of each experiment, were prepared in culture
medium and the pH was readjusted to 7.4 with HC1. TPA and ß-
phorbol were added to the culture medium from 1000-fold stock
solutions in dimethyl sulfoxide. EGF-induced down-regulation of the
EGF receptor was induced by culturing hepatocytes for 16 h with 10
fig/ml EGF. Medium containing unlabeled EGF was then removed and
5907
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