The behavior of pure cultures of nitrifying microorganisms under autotrophi
c growth operating conditions was investigated and the relations between th
eir energy metabolism and their anabolism analyzed by means of metabolic ne
twork computation. The description of the metabolism of the nitrifiers is e
xtended to their energy metabolism by introducing compartmentalization (cyt
oplasmic and periplasmic sides) and studying coupling between the electron
transport chain and the proton gradient generation. The energy model of Nit
rosomonas and Nitrobacter was developed based on the oxidoreduction reactio
ns known to be involved. The electron transport chains and the associated p
roton translocation for these models are described. Several possible hypoth
eses are analyzed and discussed concerning the thermodynamic consistency of
all the oxidoreduction reactions. For Nitrosomonas, the most delicate poin
t is the second step of hydroxylamine oxidation. For Nitrobacter a new ener
gy model is proposed in which NO plays an important role as node in the dis
tribution of electrons from NO; oxidation to the membrane electron transpor
t chain. The compartmentalization enables us to consider a proton gradient
dissipation flux as the expression of the overall energy loss in metabolic
analysis (the so-called maintenance phenomena). The energy model (electron
transport chain, proton gradient) is associated with an overall description
of the metabolism of Nitrosomonas and Nitrobacter in terms of metabolic fl
ux calculation. This representation demonstrates that a maintenance in nitr
ifiers expressed as a proton leak is no higher than for other aerobes. The
yields calculated from the energy models integrated with the metabolic mode
ls of nitrifiers are consistent with the experimental yields in the literat
ure. (C) 2001 John Wiley & Sons, Inc.