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protein can be obtained relatively easily by phage display[7] or
other approaches for in vitro evolution.[8] Although peptides
have limited utility as therapeutic agents, their efficacy,
selectivity, and specificity are often better than those of
small fragments.[9] Thus, the introduction of information from
target-protein–peptide interactions in the design of high-
affinity ligands would be an effective way to compensate for
the problem originating from the small interaction surface of
compounds.
Herein, we demonstrate that the affinity of a small-
molecule inhibitor for its target protein could be successfully
improved by the generation of hybrid molecules in which non-
overlapping moieties from a competitive peptide were
introduced. In this strategy, overlapping and non-overlapping
peptide–compound pharmacophores were discriminated by
using INPHARMA. This novel approach combines the
advantages of peptides and small molecules for the develop-
ment of higher-affinity ligands by the ligand-based NMR
spectroscopic approach.
Figure 1. Structural representation of pep-10L peptide and losartan.
A) The GPVI-bound structure of pep-10L, as determined by structural
calculation by the use of transferred NOE (TrNOE) information.[13]
Only residues 4–10, which represent the minimal binding sequence for
binding to GPVI, are shown. Residue numbers and three-letter codes
are indicated. The residues important for GPVI binding are indicated
by black boxes. B) The GPVI-bound structure of losartan.[15] All carbon
atoms in losartan are numbered. The carbon atoms in the phenyl-
tetrazole (PTZ) moiety, which is important for GPVI binding, are
circled.
Our approach was applied to the interaction between the
platelet receptor glycoprotein VI (GPVI) and collagen. GPVI
is primarily responsible for initial collagen-dependent platelet
adhesion and aggregation; deficiency of GPVI causes a loss of
collagen-stimulated platelet activation.[10] Interestingly, the
absence of GPVI does not significantly prolong bleeding time,
and a recent phase I clinical study demonstrated that GPVI–
Fc dimers (Fc is the fragment crystallizable region of an
antibody) inhibited collagen-induced platelet aggregation
without affecting general hemostasis.[11] Therefore, unlike
other antiplatelet agents, which are often associated with
adverse bleeding events, an inhibitor of the GPVI–collagen
interaction would be an interesting target for a safer anti-
aggregation drug. In fact, it has been shown that the
nonpeptide angiotensin II type 1 (AT1) receptor antagonist
losartan and its metabolite also inhibit collagen-dependent
platelet activation by GPVI at the clinical dose.[12]
Recently, we used an approach based on a phage-dis-
played peptide library to create a 12 amino acid peptide, pep-
10L (H2N-YSDTDWLYFSTS-COOH), which inhibits the
GPVI–collagen interaction.[13] The structure of pep-10L in the
GPVI-bound state was determined by structural calculation
by using transferred NOE (TrNOE) information.[13] The
structure revealed that the central part of pep-10L (Asp5-
Phe9) adopts a helical conformation (Figure 1A), in which
the side chains of Trp6, Leu7, and Phe9 form a hydrophobic
cluster upon binding to GPVI.[13] By the use of STD experi-
ments and site-directed mutagenesis, the hydrophobic cluster
was identified as the key interaction site of pep-10L.[13] The
chemical-shift perturbation (CSP) of GPVI resonances upon
binding to pep-10L indicated that the peptide binds to the
proposed collagen-binding site[14] with a KD value of 5.7 ꢀ
10À5 m (see Figure S1 in the Supporting Information).
chemical structure for the interaction with GPVI (Fig-
ure 1B).[15]
To determine whether there was direct competition
between losartan and pep-10L on the GPVI surface, we
performed NMR spectroscopic competition experiments. We
1
recorded H–15N correlated spectra of 15N-labeled pep-10L
alone (see Figure S2A), in the presence of unlabeled GPVI–
Fc (see Figure S2B), and after adding losartan to the mixture
(see Figure S2C). When the pep-10L/GPVI-Fc ratio was set to
1:0.25, the signal intensity of pep-10L significantly decreased
in comparison to the free state (see Figure S2B). This
decrease in intensity reflects accelerated relaxation owing to
interaction with a larger molecule, GPVI–Fc. Upon the
subsequent addition of losartan (25-fold concentration rela-
tive to pep-10L) to the mixture of pep-10L and GPVI–Fc, the
intensity of the pep-10L resonances were restored to up to
80% of their original intensity (see Figure S2C). These results
indicated that pep-10L and losartan compete for the same
binding site in the GPVI molecule.
With the competing low-molecular-weight compound
losartan and the peptide pep-10L, we started to investigate
which parts of the compound overlap with pep-10L by using
1
the INPHARMA method (Figure 2). Two-dimensional H–
1H NOESY experiments were performed with various
mixing times (60, 100, 200, and 300 ms) with losartan
(1.4 mm), pep-10L (0.9 mm), and GPVI–Fc (25 mm) in D2O
buffer. At a mixing time of 200 ms, interligand NOE peaks
between the PTZ moiety in losartan and pep-10L were clearly
observed in the NOESY spectrum (Figure 2C). Interligand
NOE peaks were not detected in the absence of GPVI–Fc or
with the Fc portion alone, thus excluding the possibility of
a direct interaction between losartan and pep-10L or of
indirect magnetization transfer via the Fc fragment (data not
shown). In view of the competition between pep-10L and
losartan for the GPVI–Fc binding site, these NOE peaks
could be identified as protein (GPVI)-mediated
INPHARMA peaks.
Interestingly, the site seems to overlap with the binding
site of losartan, which we proposed recently on the basis of
NMR spectroscopy.[15] By using NMR spectroscopic methods
in combination with in silico tools, we found that losartan
specifically interacts with extracellular immunoglobulin (Ig)-
like domain 1 of GPVI with a KD value of 1.7 ꢀ 10À4 m, and
that the phenyltetrazole (PTZ) moiety in losartan is the key
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ꢀ 2014 Wiley-VCH Verlag GmbH & Co. KGaA, Weinheim
Angew. Chem. Int. Ed. 2014, 53, 2597 –2601