Curr. Med. Chem. – Central Nervous System Agents, 2004, 4, 000-000 1
1568-0150/04 $45.00+.00 © 2004 Bentham Science Publishers Ltd.
The Controversial Role of Adenosine A
2A
Receptor Antagonists as
Neuroprotective Agents
Patrizia Popoli
a*
, David Blum
b
, Annita Pintor
a
, Maria Teresa Tebano
a
, Claudio Frank
a
,
Marco Gianfriddo
c
, Maria Rosaria Domenici
a
, Serge N. Schiffmann
b
and Felicita Pedata
c
a
Department of Pharmacology, Istituto Superiore di Sanità, Rome (Italy),
b
Laboratory of Neurophysiology, Université
Libre de Bruxelles, 1070, Brussels, (Belgium),
c
Department of Preclinical and Clinical Pharmacology, University of
Florence (Italy)
Abstract: Adenosine is an endogenous modulator that regulates many Central Nervous System functions, and whose
effects are mediated by four G-protein coupled receptors (A
1
, A
2A
, A
2B
, A
3
). Adenosine A
2A
receptors have been recently
regarded as promising targets for the development of neuroprotective strategies. In particular, since an abnormal
glutamate outflow is thought to play a crucial role in triggering the cellular events leading to excitotoxic neuronal death,
and since A
2A
receptors positively modulate glutamate outflow, it has been supposed that their blockade could represent a
suitable approach to the treatment of neurodegenerative diseases. In agreement with this hypothesis, both the genetic
inactivation and the pharmacological blockade of A
2A
receptors proved effective in several models of brain injury. In
some studies, the neuroprotective effects of A
2A
receptor antagonists correlated well with their ability to prevent injury- or
toxin- stimulated glutamate outflow. The “beneficial” effects of A
2A
receptor antagonists at the pre-synaptic sites (namely
their ability to attenuate glutamate release), however, seem to occur only at very low doses and at certain time points after
the insult is given. Moreover, A
2A
receptor antagonists seem unable to prevent the toxic effects elicited by direct
stimulation of post-synaptic NMDA receptors.
It is concluded that, although A
2A
antagonists show clear neuroprotective effects in models of brain injury, their actual
therapeutic potential needs to be confirmed in a wider range of doses, at different stages of brain injury as well as in
models of neurodegenerative diseases, in which pre- and post-synaptic effects play different relative roles.
Keywords: Adenosine A
2A
receptor antagonists; Neuroprotective effects; Glutamate release; NMDA receptor stimulation;
Dose-dependency; Inflammatory mechanisms.
1. THE NEUROMODULATORY ROLE OF
ADENOSINE IN THE CENTRAL NERVOUS SYSTEM
(CNS)
Adenosine is an endogenous modulator which is present
virtually in all tissues and which plays a pivotal role in
intercellular communication.
In the CNS, adenosine may act as a neurotransmitter, a
neuromodulator or “a retaliatory metabolite” [1]. The main
pathways of adenosine production, release and metabolism
in the brain have been recently reviewed by Latini and
Pedata [2]. The main source of adenosine is represented by
ATP whose intracellular degradation proceeds via adenylate
kinase to ADP and AMP, and then formation of adenosine
via cytosolic 5'-nucleotidase. Although the mechanism most
commonly implicated in adenosine release is through bi-
directional nucleoside transporters which transfer adenosine
following its concentration gradient, evidence of a neuronal
Ca
2+
-dependent release of adenosine does not exclude the
existence of an excitation-coupled release mechanism. In the
last years ATP was accepted as a neurotransmitter or
modulator in the CNS [3]. Since adenosine may also derive
*Address correspondence to this author at the Department of Pharmacology
Istituto Superiore di Sanità Viale Regina Elena, 299 00161 Rome Italy; Tel:
+39-06-49902482; Fax: +39-06-49387104; E-mail: patrizia.popoli@iss.it
from released ATP, which is dephosphorilated to AMP
which on its turn is converted to adenosine by extracellular
5'-nucleotidases [4], the contribution of extracellularly
released nucleotide to extracellular adenosine remains a
matter of debate. Under conditions of hypoxia/ischemia in
which breakdown of ATP occurs and there is a mismatch
between ATP degradation and resynthesis, adenosine
intracellular concentration greatly increases, and it is likely
that under these conditions extracellular adenosine mainly
derives per se from cells.
In in vivo and in vitro studies, it was evaluated that
adenosine is present extracellularly at concentrations which
can stimulate its higher affinity A
1
and A
2A
receptors. Under
conditions of hypoxia/ischemia, adenosine rises to
concentrations which can stimulate also its lower affinity A
3
and A
2B
receptors. Under these last conditions, adenosine is
considered as an homeostatic cell signal, which tends to
maintain the resting state by counteracting uncontrolled
activation of neuronal and glial cells [5], and as a promoter
of trophic effects in both neurons and glia (reviewed in [6]).
In heart and brain, adenosine may favor tissue adaptation to
ischemic insults (reviewed in [7]). Although adenosine
effects are generally regarded as beneficial, a great
complexity arises from the different distribution and the
different functions of its receptors [8].