A Selective Tumor Microvasculature Thrombogen that
Targets a Novel Receptor Complex in the Tumor
Angiogenic Microenvironment
Amr El-Sheikh,
1
Per Borgstrom,
2
Gourab Bhattacharjee,
1
Mattias Belting,
1,3
and Thomas S. Edgington
1
1
Department of Immunology, The Scripps Research Institute;
2
Vascular Biology and Angiogenesis Program, The Sidney Kimmel Cancer
Center, La Jolla, California and
3
Department of Cell and Molecular Biology, Lund University, Lund, Sweden
Abstract
We have previously shown that part of the heparin-binding
domain of the vascular endothelial growth factor (VEGF),
designated HBDt, localizes very selectively to surfaces of the
endothelial cells of i.t blood vessels. Here, we have coupled
the HBDt to the extracellular domain of tissue factor (TFt),
to locally initiate the thrombogenic cascade. In tumor-
bearing mice, infusion of this HBDt.TFt results in rapid
occlusive thrombosis selective only for tumor microvascula-
ture with resultant infarctive destruction of tumors. We now
show that infusion of an optimal combination of this
HBDt.TFt and its requisite cofactor ( factor VIIa) in tumor
models results in significant tumor eradication. Binding
studies and confocal microscopy indicate that the target for
the HBDt.TFt seems to be a trimolecular complex of
chondroitin C sulfate proteoglycan, neuropilin-1, and VEGF
receptor-2, overexpressed together only in highly angiogenic
sites of the tumor microenvironment. The HBDt.TFt was
also colocalized with the trimolecular receptor complex in
endothelial sprouts from tumor tissues, and its binding
inhibited the growth of such sprouts. In vitro , we show that
the HBDt structure has its highest affinity for chondroitin 6
sulfate. We show the potential of this HBDt.TFt as a
candidate therapeutic and elucidate its target in vivo . (Cancer
Res 2005; 65(23): 11109-17)
Introduction
To search and discover novel targets specific for tumor vascular
endothelium is a challenging task but a critical aspect of
developing more effective tumor therapeutics. The existence of
such targets expands on our perception of the uniqueness of the
tumor microenvironment. To be an optimally effective target, it
should not only be expressed selectively by tumor microvasculature
but should also be a common feature found in a broad spectrum of
solid tumors.
We previously showed that the rather novel vascular endothelial
growth factor (VEGF) heparin-binding domain substructure
(HBDt), when expressed on the surface of a modified M13 phage,
selectively enabled accumulation in vivo on the vascular surface of
the CD31-positive endothelium of tumor vasculature (1). The
VEGF
165
isoform is known to bind to VEGF receptor-2 (VEGFR-2,
KDR), neuropilin-1 (Npn-1), and an unknown heparan sulfate (HS)
proteoglycan (2). In addition to VEGFR-2, Npn-1 is also known to
be highly expressed by endothelial and tumor cells (3) and is
known to play important roles in endothelial mitogenesis, tumor
cell migration, and invasion (4–7). The role of Npn-1 expression in
tumor angiogenesis has been observed in several types of tumors
(4, 8). Furthermore, it was reported that expression of Npn-1 in
tumors seems to correlate with a shift from benign stromal tissue
to that associated with malignancies (9).
The VEGF
165
isoform, which contains HBDt, is known to
participate in the formation of molecular complexes, containing
VEGFR-2 and Npn-1. The HBDt-encoded VEGF/exon 7 is thought
to interact with the Npn-1 b1b2 domain (10–12). The b1b2
region of Npn-1 was also analyzed and shown to be a heparin-
binding domain, which enhanced the binding of VEGF
165
to Npn-
1 (13). The size of the oligosaccharide fragment needed for
enhancement of binding was at least 20-monosaccharide unit in
length, suggesting the potential involvement of a large polysac-
charide chain, possibly a chondroitin sulfate proteoglycan
(CSPG).
We fused the HBDt structure to the extracellular domain of
human tissue factor (TFt) through a linker to explore the feasibility
of targeting the truncated extracellular domain of TFt to the tumor
microvessel plasmalemma in a mode to locally initiate the
thrombogenic cascade. Local infarctive necrosis of tissue follows
loss of local vascular perfusion; a concept conserved in evolution
from marine invertebrates to mammals.
We have now analyzed the ability of the HBDt.TFt fusion
protein to bind heparin and other carbohydrate moieties in vitro
and to endothelial cells in culture. We also investigated the effect
of HBDt.TFt addition on angiogenic sprouting ex vivo . We also
analyzed the complex dynamics of targeting of the HBDt.TFt
protein to tumor intravascular microenvironment when infused
in vivo in various murine tumor models, monitoring thrombus
formation by real-time intravital microscopy. We investigated
the effect of HBDt.TFt administration and the resultant throm-
bosis on the destruction of tumors. Finally, we confirmed docking
of HBDt.TFt in the i.t. vasculature and colocalization with
its target receptor molecules by confocal microscopy in vivo and
ex vivo . Our data show that HBDt.TFt accumulates in i.t.
vasculature preferentially, attributable to recognition of a unique
target, and docks on tumor endothelium plasmalemma, in
a manner to permit functional initiation of the coagulation
cascade. This results in vascular occlusion of tumors, resulting in
local infarctive necrosis and significantly reducing the mass of
solid tumors. We support the potential of this strategy for
therapeutic purposes and illustrate the unique biology of the
tumor microenvironment.
Requests for reprints: Thomas S. Edgington, Department of Immunology (SP258),
The Scripps Research Institute, 10550 North Torrey Pines Road, La Jolla, CA 92037.
Phone: 858-784-8225; Fax: 858-784-8480; E-mail: tse@scripps.edu.
I2005 American Association for Cancer Research.
doi:10.1158/0008-5472.CAN-05-2733
www.aacrjournals.org 11109 Cancer Res 2005; 65: (23). December 1, 2005
Research Article
Research.
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