[CANCER RESEARCH 51, 5587-5595. October 15. 1991]
Induction of Tissue-type Plasminogen Activator by Ionizing Radiation in Human
Malignant Melanoma Cells1
David A. Boothman,2 Meizhi Wang, and Sam W. Lee
Department of Radiation Oncology. Division of Cancer Biology, University of Michigan, Ann Arbor, Michigan 48109 [D. A. B., M. H'J, and Dana-Farher Cancer
Institute, Division of Cancer Genetics, Harvard University, Boston, Massachusetts 02115 [S. H'. L.]
ABSTRACT
Two differently timed extracellular and intracellular enzymatic and
mRNA peaks of tissue-type plasrninogen activator (t-PA) were induced
following ionizing radiation. The first peak appeared within 10 min
following \-irradiation but rapidly declined. The appearance of early t-
PA mRNA transcripts and enzymatic activity were not prevented by
actinomycin D treatment. In contrast, cycloheximide prevented the early,
minor enzymatic induction peak of t-PA. Stabilization of t-PA mRNA
transcripts appears to be an early initial response of human cells to
ionizing radiation, since the synthesis of new mRNA transcripts within
the first 30 min was not observed via nuclear run-on analyses.
Nearly 12 h following \-irradiation, a second major enzymatic peak
of t-PA was observed. Cycloheximide or actinomycin D treatments
blocked the later t-PA response. t-PA mRNA levels were induced > 100-
fold in 4 h by ionizing radiation as assayed via Northern or nuclear run-
on analyses. During the major induction period, t-PA mRNA transcripts
reached their maximum levels at 4-8 h, and intracellular enzyme levels
accumulated 6-8 h after X-irradiation. Unirradiated Ul-Mel cells dem
onstrated only low basal levels of t-PA mRNA and enzymatic activity.
Similar induction responses were found following UV-irradiation or 12-
O-tetradecanoyl-phorboI-13-acetate (I'M A) treatments.
Normal human fibroblast (i.e., GM 2936B, GM2907A, and IMR-90)
cells also demonstrated the induction of t-PA, although only one later
enzymatic peak was detected. The induction of t-PA mRNA levels and
intracellular and extracellular enzymatic activities for these cells were
50-fold lower than with Ul-Mel cells given equitoxic doses of X-rays.
Differential expression of t-PA in some tumor as compared to normal
tissues may be utilized in future chemotherapeutic regimens.
INTRODUCTION
Several laboratories have reported the induction of various
gene products in mammalian cells by a wide range of agents
that cause cytotoxic stress (see Refs. 1-3 for reviews). The
regulation of new gene transcripts and gene products following
various DNA insults now include: (a) DNA binding transcrip
tion factors, such as the UV-responsive element binding factors
and X-ray-induced protein 175 (XIP175 bp) (1, 4-6); (b) pro-
tooncogenes, such as c-fos (7), c-jun (8), and c-myc (9); (c)
several growth-related genes, such as PCNA (10), protein kinase
C (11), interleukin-1 (11), and p53 (12); and (d) a variety of
other genes, including proteases (13, 14), RNA polymerase-ß
(15), tumor necrosis factor (16), metallothioneins (17, 18), and
DT Diaphorase (19).' These gene products may influence later
events within damaged mammalian cells, including mutagene-
sis, carcinogenesis, metastasis, cell survival and repair, and cell
Received 2/22/91; accepted 8/8/91.
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
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' This work was supported by a grant from the American Cancer Society (PDT-
412).
'To whom requests for reprints should be addressed, at Department of
Radiation Oncology, Division of Cancer Biology, University of Michigan. 1331
East Ann Street. Rm 4131, Ann Arbor. MI 48109.
J D. A. Boothman, S. W. Lee, M. Wang, and E. N. Hughes. Isolation and
partial characterization of X-ray-inducible gene transcripts from radioresistant
human melanoma cells, in preparation, 1991. Presented at the 82nd annual
meeting of the American Association for Cancer Research, abstract 1671, p. 281.
Houston, TX, 1991.
death (i.e.. apoptosis) (1,3, 20-22).
Using differential hybridization, we have recently isolated 12
cDNA4 clones which were dramatically induced by ionizing
radiation; the results and specific details of the differential
hybridization screen will be described elsewhere.4 After the
clones were isolated, we partially sequenced the cDNA inserts,
and their identities (if known) were elucidated via GenBank
sequence analyses.1 One of the cDNA clones isolated contained
>95% DNA sequence homology to human tissue-type plasrnin
ogen activator.
t-PA is a secreted serine glycoprotease of approximately M,
70,000, which is produced by various different cell types and
tissues and by many human tumor cells in culture (23-26).
Both t-PA and its related serine protease, u-PA, participate in
the regulation of a multitude of cellular processes leading to
the formation of active plasmin, which then initiates a number
of cascade reactions. These proteases are therefore thought to
be important in circulatory fibrinolytic activity and may act in
concert to initiate prohormone processing, inflammation, re
production, and cell migration, including an involvement in
tumor cell formation and metastasis (see Ref. 23 for review), t-
PA has been localized to chromosome 8, and its cDNA and
complete genomic sequence have been reported (27-29).
Plasminogen activators are regulated at the enzymatic level
by various external stimuli. Increased levels of u-PA and/or t-
PA can be induced by hormones or hormone-like agents in
distinct tissue-specific manners (25, 30, 31). The induction of
t-PA has been associated with the intracellular induction of
protein kinase C (32). Increased secretion of u-PA enzymatic
activity has been reported with various mammalian cells in
response to exposure to a limited number of alkylating agents
(13, 14, 33). The induction of t-PA by ionizing radiation has
not been previously demonstrated, nor has the induction/sta
bilization of t-PA been investigated at the level of transcription
following any DNA-damaging agent.
The purpose of the present study was to further investigate
the induction of t-PA by ionizing radiation. The temporal
induction of enzymatic extra- and intra-cellular levels of t-PA
and the regulation of its mRNA were followed after exposure
of confluence-arrested Ul-Mel cells to various doses of ionizing
radiation and after exposure to other DNA-damaging agents.
We also compared the induction of t-PA in Ul-Mel cells to
that induced by density-arrested normal human fibroblasts (i.e.,
GM2936B, GM2907A, and IMR-90).
4 The abbreviations used are: cDNA, complementary DNA; Ul-Mel, radiore
sistant human malignant melanoma; t-PA, tissue-type plasminogen activator; u-
PA, urokinase-type plasminogen activator; PMA, 12-O-tetradecanoyl-phorbol-
13-acetate:f!-PKC, beta-protein kinase C; PLDR, potentially lethal DNA damage
repair.
5 N. A. Fukunaga, R. A. Schea, H. L. Burrows, and D. A. Boothman. Enhanced
induction of tissue-type plasminogen activator in human cancer-prone as opposed
to normal cells, in preparation. 1992. Presented at the Miami Winter Symposium,
Miami, FL, Feb., 1991.
5587
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