[CANCER RESEARCH 52. 3011-3014, June I, 1992]
Stimulation of Anchorage-independent Cell Growth by Endothelin in NRK 49F
Cells1
Masatoshi Kusuhara,2 Ken Yamaguchi, Masaru Kuranami, Ai Suzaki, Shiro Ishikawa, Hanlim Moon, Isamu Adachi,
Shingo Hori, and Shunnosuke Handa
Growth Factor Division, National Cancer Center Research Institute [M. Kus, K. Y., M. Kur, A. S., S. I., H. M.j and Department of Medicine, National Cancer Center
Hospital [I. A.], Tsukiji 5-1-1, Chuo-ku, Tokyo 104; and Department of Medicine fS. Ho., S. Ha.], Keio University School of Medicine, Shinanomachi 35, Shinjuku-
ku, Tokyo 160, Japan
ABSTRACT
Endothelin (ET) is a vasoconstrictor peptide originally isolated from
vascular endothelial cells. Recent studies have revealed that ET has many
biological functions including growth factor-like activity. The present
study aims to clarify whether ET-1 possesses the ability to stimulate
anchorage-independent cellular growth, an indicator of factors with trans
forming activity.
We found that NRK 49F cells possess a large number of high-affinity
ET-1 receptors; labeled '"I-ET-1 binding was displaced by unlabeled
ET-2 in a similar dose response, but in the case of ET-3, 100-fold more
was required. Specific '"I-ET-3 binding was undetectable in NRK 49F
cells, indicating that ET receptors in NRK 49F cells are 11-1112
selective.
NRK 49F is a cell line which is most commonly used to assay for
anchorage-independent cellular growth. Therefore, we explored whether
ETs promote anchorage-independent cellular growth in this cell line. II-
1 and ET-2 stimulated NRK colony formation dose dependent!)' in the
presence of 1 MMepidermal growth factor (EGF). In contrast, ET-3 did
not have colony-stimulating ability. In the presence of EGF, the maximal
effect of ET-1 was approximately 90% of that of transforming growth
factor-/?. Moreover, in the presence of maximal stimulating concentra
tions of EGF and transforming growth factor-/?, ET-1 additionally induced
colony formation. These results indicate that ET-1 and -2 possess trans
forming growth factor-like activity for NRK 49F cells. Since ET-1 and
-2 increased intracellular calcium levels, this ion may participate in signal
transduction pathways by which ET-1 and -2 promote colony formation.
INTRODUCTION
ET-13 was isolated originally as a factor possessing vasocon-
strictive activity from culture media conditioned by primary-
cultured endothelial cells (1). Three types of ET, ET-1, ET-2,
and ET-3, are now known to compose the ET family, and their
production and secretion have been demonstrated in many
tissues (2). Recent research has revealed that ET-1 also has a
wide range of pharmacological effects in tissues other than
blood vessels, such as an inotropic effect on cardiac myocytes
(3), a neurotransmitter effect on the central nerve system (4),
and modulation of endocrine secretions (5-8).
Several studies, including ours, demonstrated that ET-1 stim
ulates DNA synthesis in various types of cultured cells, includ
ing Swiss 3T3 (9, 10), vascular smooth muscle (11, 12), and
Received 6/19/91; accepted 3/20/92.
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.
1This investigation was supported in part by a research grant from the Princess
Takamatsu Cancer Research Fund, by a Grant-in-Aid from the Ministry of Health
and Welfare for the Comprehensive 10-Year Strategy of Cancer Control, and by
Grants-in-Aid for Cancer Research (1-5 and 2-9) from the Ministry of Health
and Welfare.
2To whom requests for reprints should be addressed, at Growth Factor
Division, National Cancer Center Research Institute, Tsukiji 5-1-1, Chuo-ku,
Tokyo 104, Japan.
'The abbreviations used are: ET, endothelin; DMEM, Dulbecco's modified
Eagle medium; CS, calf serum; TGF, transforming growth factor; EGF, epidermal
growth factor; [Ca2*]¡, intracellular calcium ion concentration; A'd, dissociation
constant.
renal mesangial (13). However, how ET-1 influences cellular
growth is poorly understood. Our recent observations that
cancer cells frequently produce and secrete ET-1 and that
primary-cultured human fibroblasts possess ET-1 receptors sug
gest that ET-1 produced by cancer cells could act as a paracrine
growth factor on mesenchymal cells (14).
In the present study, we explored whether ET-1 promotes
anchorage-independent cell growth. For these experiments,
NRK 49F was selected as an indicator cell line, which is most
commonly used to assay for anchorage-independent cell growth
(15, 16).
MATERIALS AND METHODS
Cell Culture and Materials. NRK 49F cells were provided by the
Japanese Cancer Research Resources Bank (Tokyo, Japan). They were
grown in DMEM (Nissui Seiyaku, Tokyo, Japan) containing 5% CS
(GIBCO Laboratories, Grand Island, NY), 100 Mgof streptomycin/ml,
and 100 units of penicillin G/ml at 37°Cin a fully humidified atmos
phere of 5% COz in air. Confluent cells were further cultured in DMEM
containing 1% CS for 24 h to minimize the effect of serum.
Synthetic ET-1, -2, and -3 were purchased from the Peptide Institute
(Osaka, Japan); recombinant TGF-/3 and human EGF from Earth
Chemical Co., Ltd. (Hyogo, Japan); bovine serum albumin (Conn
fraction V) from Dai-ichi Pure Chemicals Co., Ltd. (Osaka, Japan);
'"I-ET-1 (specific activity, 74 TBq/mmol) and '"I-ET-3 (specific activ
ity, 74 TBq/mmol) from Amersham International (Buckinghamshire,
United Kingdom); and Fura-2 acetoxymethyl ester from Molecular
Probes, Inc. (Eugene, OR).
Binding Assay. The ET-1 binding assay was performed as previously
described, with slight modification (17). NRK 49F cells were grown to
confluence in a 48-well plate and further cultured for 24 h in DMEM
containing 1% CS. Hanks' balanced salt solution containing 20 m\i N-
(2-hydroxyethyl)piperazine-yV'-(2-ethanesulfonic acid) and 0.1 % bovine
serum albumin (pH 7.4) was used as assay buffer. After these cells were
washed, they were incubated at 37°Cfor 60 min with 0.12 nM '"I-ET-
1 plus various concentrations of unlabeled ET-1, -2, and -3. They were
washed three times and solubilized with l N NaOH. Cell-bound radio
activity was determined in a gamma spectrometer. Cell numbers were
estimated by counting other wells which were treated in the same
manner as those of the binding studies. Specific binding was calculated
as total binding minus binding in the presence of 1 pM unlabeled
ET-1.
Since specific '"l-ET-3 binding could not be detected under the
above conditions, the binding assay was done by using a larger number
of NRK 49F cells. NRK 49F cells in 60-mm dishes were incubated
with 0.3 nM '"I-ET-3 in the presence or absence of a 5000-fold excess
of unlabeled ET-3.
Soft Agar Colony Formation Assay. To assess anchorage-independent
cell growth, a soft agar colony formation assay using NRK 49F cells
was performed as previously described, with slight modification (18).
NRK 49F cells (20,000/dish) were suspended in 0.9 ml of culture
medium containing 0.3% agar and spread over a prehardened feeder
layer comprising 0.9 ml of the same medium, but containing 0.5% agar,
in a 35-mm Petri dish. Test materials dissolved in 0.2 ml of culture
medium were placed on the top of the prehardened agar layers. After
14 days of incubation at 37°C in a humidified CO2 incubator, colonies
3011
on July 15, 2015. © 1992 American Association for Cancer Research. cancerres.aacrjournals.org Downloaded from