Current Cancer Drug Targets, 2005, 5, 285-298 285
Matrix Metalloproteinase Inhibitors as Anticancer Therapeutics
F. Mannello
*,1
, G. Tonti
1
and S. Papa
2
1
Institute of Histology and Laboratory Analysis;
2
Institute of Morphological Sciences; Faculty of Sciences MM.
FF. NN., University of Urbino “Carlo Bo”, 61029 Urbino, Italy
Abstract: Matrix metalloproteinases (MMPs), also designated as matrixins, play a central role in many
biological processes and are involved both in physiologic cellular processes and in pathologic situations
such as tumor growth, invasion and metastasis. For more than 30 years MMPs have been considered as
promising targets for cancer therapy and a number of different synthetic and natural MMP inhibitors have been
identified as cytostatic and anti-angiogenic agents and have begun clinical testing in view of their specific
implication in malignant tissues. Although preclinical studies were so compelling to encourage several
clinical trials, the past years have seen a consistent number of disappointments and limited success. The critical
examination of previous studies shed light on new information about the cellular source, substrates and mode of
action of MMPs, focusing the attention of future research on the identification of specific MMP targets in
tumors at different stage of tumor progression, both in order to improve efficacy and to reduce the side effect
profile. In this review we discuss the current view on the feasibility of MMPs as target for therapeutic
intervention in cancer, taking into account that the perspective may be of great value for molecular medicine for
the twenty-first century, providing intriguing information about the MMPs as mediators in biology and
pathology, and as targets for disease therapies.
Keywords: Matrix metalloproteinases, Gelatinases, Cancer, Protease Inhibitors, Therapeutic strategies, Clinical trials.
INTRODUCTION constitute the zinc binding site and a “methionin-turn” motif
that lies beneath the active site zinc ion [3]. The ion-binding
motif reads HEBXHXHBGBXHZ, where histidine (H),
glutamic acid (E) and glycine (G) are invariant, B is a bulky
hydrophobic residue, X is a variable residue and Z is a
family-specific amino acid (serine in MMPs). All MMPs
have an N-terminal hydrophobic signal sequence
(predomain) which leads their synthesis to the endoplasmic
reticulum and their secretion into the extracellular space; this
amino acid stretch is thereafter removed. This predomain is
followed by a 77-87 amino acid-long prodomain that
constitutes the N-terminus of the secreted enzyme and
maintains it in its latent form until its removal or
disruption. The prodomain keeps the enzyme inactive
through a mechanism identified as “ cysteine switch” where
the unpaired cysteine in the highly conserved “Pro-Arg-Cys-
Gly-X-Pro-Asp” sequence forms a bridge with the catalytic
zinc, thus preventing enzymatic activity. The enzyme
acquires total proteolytic capacity when the prodomain
becomes chemically removed by cleavage [1]. The active site
is of great importance: it specifically binds to selective
substrates by means of its active site cleft, through specificity
sub-site pockets that bind amino acids adjacent to the
scissile peptide bond, and through secondary substrate-
binding exosites located outside the active site [5]. These
domains represent the minimal structure of MMPs found in
MMP-7 (matrilysin) and MMP-26 (endometase/matrylisin-
2) which lack any other domain. All the other MMPs have a
hinge region varying in length and composition which also
influences substrate specificity [6] , and a four-blade β-
propeller structure representing the hemopexin/vitronectin-
like domain [7]. Two metalloproteinases, Gelatinase A and
B, are further characterized by the presence of three head-to-
tail cystein-rich repeats within the catalytic domain. This
structure resembles the collagen-binding type II repeats of
fibronectin and is necessary for the binding and cleavaging
Extracellular proteinases are involved in a wide variety of
developmental and disease-associated processes proving to
be key protagonists in many physiological and pathological
mechanisms. The first observation made 40 years ago of an
activity during metamorphosis in tadpole tails, able to
degrade interstitial collagen, opened a new field of
unexpected biomedical research and the way to study what
has now become the matrix metalloproteinase (MMP) family
[1]. These endopeptidases belong to the wider metzincin
group, which in turn constitutes one of several
metalloendopeptidase families [2-4]; all the metzincins are
zinc-dependent proteinases and according to their structural
characteristics are subdivided into serralysins, adamalysins,
astacins and matrixins [3]. As our knowledge of the MMP
family continues to grow, we are learning that their activities
are not limited to matrix degradation [1]. MMPs seem to
touch almost every aspect of mammalian biology and,
through more than 1,000 new publications each year, certain
exciting discoveries solidified the field, solving previous
puzzles and pointing the research in new directions (starting
from biochemical studies, discussing the flurry of activity
that involved the gene regulation through molecular biology
approaches, and finally focusing the interest to MMPs as
therapeutic targets in several diseases).
STRUCTURAL AND FUNCTIONAL COMPLEXITY
OF MMPs
All MMPs share a conserved structural topology, which
consists of a catalytic domain containing three histidines that
*Address correspondence to this author at the Istituto di Istologia ed
Analisi di Laboratorio, Facoltà di Scienze MFN, Via E. Zeppi, Università
degli Studi di Urbino “Carlo Bo”, 61029 Urbino (PU), Italy; Tel: +39-
0722-351479; Fax: +39-0722-322370; E-mail: f.mannello@uniurb.it
1568-0096/05 $50.00+.00 © 2005 Bentham Science Publishers Ltd.