Journal of Computer-Aided Molecular Design, 11 (1997) 153–162 153
*To whom correspondence should be addressed.
ESCOM
J-CAMD 388
Role of the tautomerism of 2-azaadenine and 2-azahypoxanthine
in substrate recognition by xanthine oxidase
Begoña Hernández
a
, Modesto Orozco
a
and Francisco J. Luque
b,
*
a
Department of Biochemistry and Molecular Biology, Faculty of Chemistry, University of Barcelona,
Martí i Franquès 1, E-08028 Barcelona, Spain
b
Department of Pharmacy, Section Physical Chemistry, Faculty of Pharmacy, University of Barcelona,
Avinguda Diagonal s/n, E-08028 Barcelona, Spain
Received 30 August 1996
Accepted 6 December 1996
Keywords: Xanthine oxidase; Molecular recognition; Tautomerism; Quantum mechanics; Xanthine
Summary
The tautomerism of 2-azaadenine and 2-hypoxanthine has been examined in the gas phase and in
aqueous solution. The tautomerism in the gas phase has been studied by means of semiempirical and
ab initio quantum-mechanical computations, as well as density-functional calculations. The influence
of the aqueous solvent on the relative stability between tautomers has been estimated from self-consist-
ent reaction field calculations performedwith different high-level continuum models. The results provide
a detailed picture of the tautomeric preference for these purine bases. The importance of tautomerism
in the substrate recognition by xanthine oxidase is discussed. Finally, the rate of oxidation of 2-aza-
adenine and 2-hypoxanthine by xanthine oxidase is discussed in terms of the recognition model at the
enzyme active site.
Introduction
The degradation metabolism of purines is mainly con-
trolled by two enzymes: adenosine deaminase (EC 3.5.4.4)
and xanthine oxidase (xanthine:O
2
oxidoreductase; EC
1.2.3.2) [1]. The former enzyme deaminates adenosine
yielding inosine, which is subsequently decomposed into
the ribose and the purine base, hypoxanthine (1,7-dihydro-
6H-purin-6-one). Oxidation at position 2 of hypoxanthine
by xanthine oxidase yields xanthine (1,7-dihydro-6H-
purin-2,6-dione), which is also an intermediate product in
the catabolism of guanine. Indeed, this enzyme catalyzes
oxidation at position 8 of xanthine leading to uric acid,
the final degradation product of purines in humans.
Xanthine oxidase is a very important pharmacological
target, since a malfunctioning of this enzyme increases the
content of uric acid and eventually leads to the deposition
of sodium hydrogen urate monohydrate crystals in joints.
This gives rise to a painful disease known as gout, which
is clinically treated by the antihyperuricemic drug allo-
purinol [2] (pyrazolo[3,4-d]pyrimidin-6-one), a drug struc-
turally related to hypoxanthine. Allopurinol is converted
by xanthine oxidase into alloxanthine (pyrazolo[3,4-d]-
pyrimidin-2,6-one). This latter compound inactivates the
enzyme [3], thus inhibiting the formation of uric acid. The
treatment of gout with allopurinol leads to the excretion
of purines mainly as hypoxanthine and xanthine.
Owing to the biochemical and pharmacological rel-
evance of xanthine oxidase, a great deal of research has
focused on the structural and kinetic aspects of the enzy-
matic mechanism [4]. Xanthine oxidase and the related
xanthine dehydrogenase (xanthine:NAD
+
oxidoreductase;
EC 1.2.1.37) both comprise two equivalent, independent
subunits [4], each containing one atom of molybdenum,
one molecule of flavin adenine dinucleotide and two
distinct iron-sulfur clusters. There are two spatially separ-
ated binding sites for the reducing (hypoxanthine, xan-
thine) and oxidizing (O
2
, NAD
+
) substrates. The enzyme
follows a two-site ping-pong mechanism [4,5], in which all
the prosthetic groups participate in catalysis forming an
electron transport chain connecting the two binding sites.
The basic catalytic unit consists of a MoOS moiety [4c,d].
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