[CANCER RESEARCH 49, 6123-6129, November 1, 1989]
Monocyte Killing of Human Squamous Epithelial Cells: Role for Thrombospondin1
Bruce L. Riser, Rajorshi Mitra, Debra Perry, Vishva Dixit, and James Varani2
Department of Pathology, University of Michigan Medical School, Ann Arbor, Michigan 48109
ABSTRACT
Human peripheral blood monocytes maintained in culture for 18 h
were examined for killing of normal human keratinocytes and squamous
carcinoma cells. Keratinocytes grown under conditions which maintain
the undifferentiated state were highly sensitive to killing but these cells
became resistant to killing after induction of differentiation. A line of
squamous carcinoma cells obtained from an undifferentiated tumor (des
ignated as UM-SCC-HB) was sensitive to killing while a second line
obtained from a more well-differentiated tumor (designated as UM-SCC-
22B) was resistant. Several observations suggested that interaction of
monocytes with the squamous epithelial cells was mediated, in part,
through thrombospondin (TSP). Monocytes synthesized TSP and were
positive by immunofluorescence for surface TSP. The normal and malig
nant squamous epithelial cells also expressed surface TSP as well as
unoccupied TSP receptors and our previous studies have shown that both
TSP and its receptor are much more prominently displayed on the
undifferentiated cells than on the differentiated cells. A series of anti-
TSP monoclonal antibodies inhibited killing. These included an antibody
directed against the M, 25,000 NH2-terminal region of the molecule
which has heparin-binding activity and three antibodies the epitopes of
which lie within the M, 140,000 non-heparin-binding fragment of TSP.
High concentrations of exogenously added TSP as well as the recombi
nant form of the heparin-binding domain from the TSP molecule also
partially inhibited killing while laminili and fibronectin were ineffective.
Taken together, these data suggest that TSP and TSP receptors on
monocytes and squamous epithelial cells play a role in monocyte-mediated
killing of the squamous epithelial cells.
INTRODUCTION
Extracellular matrix molecules are known to mediate cell-
substrate adhesion and a possible role for these molecules as
mediators of cell-cell interactions has also recently been sug
gested. Studies have shown a role for laminin in the recognition
of murine tumor cells by natural cell-mediated cytotoxic cells.
NK3 cells express laminin-like molecules on their surface (1-4)
and treatment of NK cells with antibodies to laminin inhibits
their ability to lyse target cells (1). Among murine tumor cell
lines there is a direct relationship between laminin receptor
expression and sensitivity to NK-mediated killing (5-7). Fur
ther, the addition of laminin to NK cytotoxicity assays reduces
killing of laminin receptor-positive cells (5, 6). A role for
laminin in monocyte-mediated killing of tumor cells has also
been suggested (8).
Extracellular matrix components other than laminin may
also participate in cell-cell interactions. TSP released from the
a. granules of activated platelets is thought to participate in the
secondary phase of platelet aggregation (9). Platelet-monocyte
aggregation may also be mediated by TSP. Silverstein and
Nachman (10) showed that monocytes bound TSP in a receptor-
like manner and that TSP on the surface of activated platelets
Received 11/28/88; revised 4/3/89. 8/1/89; accepted 8/7/89.
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 study was supported in part by American Cancer Society Grants IM-
432 and PDT-324.
2To whom requests for reprints should be addressed, at Department of
Pathology. University of Michigan Medical School, 1301 Catherine Road, Box
0602, Ann Arbor. MI 48109.
3 The abbreviations used are: NK, natural killer; TSP, thrombospondin; KGM,
keratinocyte growth medium; ELISA, enzyme-linked immunosorbent assay.
could bind to monocytes through this receptor. Several different
tumor cell types were subsequently reported to express a similar
receptor (11); thus it could be postulated that tumor cell-platelet
interactions may also be mediated through this mechanism. In
addition to exhibiting surface receptors for TSP, monocytes
also synthesize TSP ( 12). It is possible, therefore, that monocyte
interactions with cells other than platelets may also be mediated
by TSP. The present study suggests a role for TSP in the killing
of human squamous epithelial cells by peripheral blood mono
cytes.
MATERIALS AND METHODS
Cells. Two human squamous carcinoma cell lines (designated UM-
SCC-11B and UM-SCC-22B) were used as targets for monocyte-me
diated killing in this study. The isolation and characterization of these
lines have been described previously (13). The tumor cell lines were
grown in Eagle's minimal essential medium supplemented with non-
essential amino acids, 15% fetal bovine serum, 100 units/ml of penicil
lin, and 100 /jg/ml of streptomycin. The cells were grown at 37°Cand
5% CO2 and subcultured by trypsinization as required. In certain
experiments normal human epidermal keratinocytes were used in place
of the squamous carcinoma cells. These cells were grown in KGM
(Clonetics, San Diego, CA). This is a serum-free, low Ca2+ (0.3 ITIM)
culture medium containing epidermal growth factor, insulin, and pitui
tary extract. The keratinocytes were grown at 37°Cand 5% CO2.
Previous studies have shown that keratinocytes maintained under these
conditions remain in an undifferentiated state for several passages (14,
15). To induce differentiation, the keratinocytes were incubated in
KGM supplemented with 1.4ITIMCa2+for 2 days. In other experiments,
K562 lymphoblastoid cells were used as targets. These cells were grown
in suspension culture (37°Cand 5% CO2) using RPMI 1640 supple
mented with 10% fetal bovine serum and antibiotics as the culture
medium.
Monocytes. Human peripheral blood monocytes were isolated from
nonanticoagulated blood (usually 100 ml) that had been defibrinated
by shaking for 15 min in a 125-ml Ehrlenmeyer flask containing
approximately 90 sterile glass beads (5mm diameter). The defibrinated
blood was diluted 1:1 with RPMI 1640, layered onto Ficoll-Hypaque
(Pharmacia, Piscataway, NJ), and centrifuged at 600 x g for 25 min at
20°C.After centrifugation, the mononuclear cell layer was removed,
washed once, and resuspended in 7 ml of RPMI 1640. The cell suspen
sion was then divided into two aliquots, layered on Sepracell-MN
(Sepratech Corporation, Oklahoma City, OK), and centrifuged at 600
x g for 30 min. The monocyte layer was then removed, washed twice,
and resuspended in RPMI 1640. Cells (2.5 x IO5) in 200 n\ of RPMI
1640 were added per well to 96-well plates for use in cytotoxicity assays.
After 2 h at 37°Cand 5% CO2, the nonadherent cells were removed by
washing three times with warm RPMI 1640. Fresh RPMI 1640 with
or without 10% fetal bovine serum was then added and the monolayer
cultures were maintained at 37°Cand 5% CO2 until the time of use
(usually 18 h later). These populations were normally greater than 96%
monocytes as determined by morphology (Wright-Giemsa stain) and
the production of a-naphthol esterase (Sigma kit No. 90-AL; Sigma
Chemical Co., St. Louis, MO).
Lymphocytes. Peripheral blood lymphocytes were obtained from the
same blood as monocytes. After separation of the mononuclear cell
layer by centrifugation through Sepracell-MN, the lymphocyte layer
was removed and washed twice. The cells were finally resuspended in
5 ml of RPMI 1640 and counted in the presence of trypan blue to
determine viable cells. After counting, the cells were seeded into wells
of a 96-well plate at 2 x IO5cells/well (100 n\) and incubated overnight.
6123
Research.
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