Introduction
Bacterial N oxide metabolism is
related to cellular bioenergetics and processes of nitrogen
assimilation.
The interest in nitric oxide (NO) centers around the dissimilatory
transformation
of nitrate, better known as the denitrification process.
Denitrification
[described as phenomenon more than 100 years ago (Gayon and Dupetit
1886)],
is a distinctive mode of respiration that satisfies the bioenergetic
needs
of a great variety of bacteria by transforming oxyanions of nitrogen to
N2, mainly under conditions of reduced oxygen
tension
or strict anaerobiosis. The reaction reverses nitrogen fixation in the
biogeochemical N cycle sustained by prokaryotes. Denitrification is
controlled
by the metalloenzymes nitrate reductase, nitrite reductase, nitric
oxide
reductase, and nitrous oxide (N2O) reductase and
involves
the corresponding enzyme substrates. The same magnitude of nitrogen
fixed
yearly by biological and abiological processes [combined estimates vary
between 254 to 406 million tons N (Jenkinson 1990)] has to be returned
to N2 by denitrification to close the N
cycle.
However, because of the large anthropogenic contribution, fixation and
denitrification are not balanced anymore as evident from the steady
increase
of nitrate in the environment. A second concern focussing around NO and
the conditions of its microbial formation, is the nitrosation of
secondary
amines in the etiology of certain types of cancer. Since the discovery
of NO in 1987 as a vasodilatory messenger (Ignarro et al. 1987; Palmer
et al. 1987), the biomedical community is astounded by the diverse
roles
of NO in cellular communication including the central and peripheral
nervous
system, and in host defense mechanisms of eukaryotes (for reviews see
Marletta
et al. 1990; Moncada 1992; Nathan 1992; Traylor and Sharma 1992;
Edelman
and Gaily 1992). Yet NO is not an obscure chemical and certainly no
newcomer
to the life sciences, as often stated in hyperbole. Early in evolution
NO took its role as a central player in bacterial bioenergetics and in
the global N cycle vital to all organisms. The chemistry of NO
in
biological systems and that of the nitroxyl anion (NO-) and
nitrosonium cation (NO+) has been reviewed briefly (Stamler
et al. 1992c; see also comment by Bonner and Hughes 1993). Another
remarkable
finding is the formation of N2O from nitrite, sometimes accompanied by
NO production, by the fungus imperfectus Fusarium oxysporum and
telemorphic and anamorphic relatives (Shoun et al. 1992). These fungi
synthesize
a special cytochrome P-450 induced only in the presence of nitrite
(Shoun
and Tanimoto 1991), which has been shown to have NO reductase activity
(Nakahara et al. 1993). The existence of such a hemoprotein is of
interest
in the context that the cytokine (interferon-g
and lipopolysaccharide)-inducible form of NO synthase from macrophages
is a cytochrome P-450 (White and Marletta 1992); also hepatic
cytochrome
P-450 monooxygenases are able to convert the NG-hydroxy-activated
form of L-arginine to NO (Boucher et al. 1992). In another fungus, the
slime mold Dictyostelium discoideum, NO alters the cellular
aggregation
behavior via ADP-ribosylation of a cytoplasmic 41-kDa protein (Tao et
al.
1992). Principles of bacterial NO metabolism were spelled out nearly 40
years ago. NO was found as a product of denitrification from a clay
loam
microcosm with 15N nitrate as tracer
(Wijler
and Delwiche 1954). The study of nitrite utilization with Thiobacillus
denitrificans established that NO was consumed and produced by a
defined
axenic culture (Baalsrud and Baalsrud 1954). NO was given status of an
intermediate in bacterial denitrification, first as the result of
studies
with intact cells (Iwasaki et al. 1956; Matsubara and Mori 1968) and
later
from recognition that cell-free extracts of "Pseudomonas
denitrificans"
reduce exogenous NO (Miyata et al. 1969). Around that time Pseudomonas
stutzeri ZoBell (formerly
P. perfectomarina) was introduced
to denitrification research and a pathway identical to that of Mori and
coworkers was formulated (Payne et al. 1971):
NO3-®
NO2-®
NO
®
N2O
® N2
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