Current Topics in Medicinal Chemistry, 2007, 7, 000-000 1
1568-0266/07 $50.00+.00 © 2007 Bentham Science Publishers Ltd.
Snake Venom Phospholipase A
2
Inhibitors: Medicinal Chemistry and
Therapeutic Potential
Silvana Marcussi
a,b
, Carolina D. Sant’Ana
a
, Clayton Z. Oliveira
a
, Aristides Quintero Rueda
a,c
,
Danilo L. Menaldo
a
, Rene O. Beleboni
d
, Rodrigo G. Stabeli
e
, José R. Giglio
b
, Marcos R. M. Fontes
f
and Andreimar M. Soares
a,
*
a
Departamento de Análises Clínicas, Toxicológicas e Bromatológicas, Faculdade de Ciências Farmacêuticas de
Ribeirão Preto, FCFRP, Universidade de São Paulo, USP-RP, Ribeirão Preto-SP, Brazil;
b
Departamento de
Bioquímica e Imunologia, Faculdade de Ciências Médicas de Ribeirão Preto, FMRP, Universidade de São Paulo, USP-
RP, Ribeirão Preto-SP, Brazil;
c
Departamento de Química, Faculdade de Ciencias Naturaes e Exactas, Universidade
Autónoma de Chiriquí, UNACHI, Chiriquí, Panamá;
d
Departamento de Biotecnologia, Universidade de Ribeirão Preto,
UNAERP-SP, Ribeirão Preto, Brazil;
e
Instituto de Pesquisas em Patologias Tropical, IPEPATRO, Universidade
Federal de Rondônia, UNIR, Rondônia-RO, Brazil;
f
Departamento de Física e Biofísica, Instituto de Biociências,
Universidade Estadual Paulista, UNESP, Botucatu-SP, Brazil.
Abstract: Phospholipases A
2
(PLA
2
s) are commonly found in snake venoms from Viperidae, Hydrophidae and Elaphidae
families and have been extensively studied due to their pharmacological and physiopathological effects in living
organisms. This article reports a review on natural and artificial inhibitors of enzymatic, toxic and pharmacological effects
induced by snake venom PLA
2
s. These inhibitors act on PLA
2
s through different mechanisms, most of them still not
completely understood, including binding to specific domains, denaturation, modification of specific amino acid residues
and others. Several substances have been evaluated regarding their effects against snake venoms and isolated toxins,
including plant extracts and compounds from marine animals, mammals and snakes serum plasma, in addition to poly or
monoclonal antibodies and several synthetic molecules. Research involving these inhibitors may be useful to understand
the mechanism of action of PLA
2
s and their role in envenomations caused by snake bite. Furthermore, the
biotechnological potential of PLA
2
inhibitors may provide therapeutic molecular models with antiophidian activity to
supplement the conventional serum therapy against these multifunctional enzymes.
Keywords: Phospholipases A
2
, phospholipase A
2
inhibitors, natural and artificial inhibitors, snake venoms.
1. INTRODUCTION
Global health statistics for incidence of snake bite
envenomations and their severity remain unknown or misun-
derstood. In spite of the lack of data, a global estimation of
the number of ophidian accidents reaches one million cases
per year, accounting for 20,000 deaths, especially along rural
areas in Asia, South America and Africa. In addition to
mortality, these envenomations are also a public health
concern as a result of the chronic morbidity associated with
them (e.g. amputations, deformations and renal failure),
which causes significant social and economic impact [1, 2].
Snake venoms induce shock, proteolysis, blood clotting,
release of bioactive substances such as histamine and
bradykinin, hemorrhage, necrosis and several other effects
[3]. Necrosis may result from a direct action of myotoxins or
myotoxic PLA
2
s on muscle cell plasma memb-ranes, or
indirectly, as a consequence of blood vessel dege-neration
and schemia caused by hemorrhagins. Disorga-nization of
membrane phospholipid components promotes release of
intracellular creatine kinase, which may be used as a
biomarker for myotoxic activity evaluation [4, 5].
The high medical-scientific concern evoked by the
involvement of these proteins in different physiopathological
processes promoted an increasing search for natural or
*Address correspondence to this author at the FCFRP-USP, Ribeirão Preto-
SP, Brasil; Fax: 55-16-3602-4725; E-mail: andreims@fcfrp.usp.br
artificial inhibitors, aiming at PLA
2
s neutralization, and a
better understanding of their mechanism of action and their
structure-function relationship. Different venom toxins can
be recognized and inhibited by poly or monoclonal
antibodies, as well as by some diversified agents including
chemicals (EDTA, BPB, and others), as well as animal and
vegetal compounds [6-9], such as heparin.
Low molecular weight heparin, an anionic natural
polysaccharide, showed to be able to inhibit, at least in part,
the myotoxic, cytotoxic and edema inducing effects of B.
moojeni MjTX-II and B. neuwiedi BnSP-7 venoms and able
to abolish the neuromuscular effect of the latter. These data
confirmed the hypothesis that the 115-129 C-terminal region
of B. asper myotoxin II, named heparin binding region,
would be responsible for PLA
2
cytotoxicity and at least in
part for its myotoxicity, since this polyanion interacts with
this region of the enzyme [9-13].
Some PLA
2
inhibitors may be found in different orga-
nisms. Manoalide (A) is a non-steroidal sesquiterpenoid
from the marine sponge Luffariella variabilis, while
manoalide (B) was artificially synthesized based on its
natural analogue. These terpenoids compounds promote
irreversible inhibitory effects upon several PLA
2
s from
snakes, bees and mammals [14, 15]
Aside from traditional knowledge or the popular use,
several scientific evidences have pointed to the efficacy of
crude extracts or fractions from plants, such as Eclipta