Fig. 5. Regulation of gene expression by redox-sensitive Arc and Fnr pathways.
Under aerobic conditions, ArcB and ArcA are sequentially activated by
phosphorylation. ArcA negatively regulates the transcription of the
cyoABCDE operon, which encodes cytochrome bo oxidase (high
Vmax, low oxygen affinity) and positively regulates
cydAB encoding cytochrome bd oxidase (low
Vmax, high oxygen affinity). Fnr is inactive under aerobic
conditions, but at low oxygen it undergoes a conformational change, probably
mediated by reduction of ferric to ferrous iron at an iron-sulphur center.
Conformationally activated Fnr binds DNA at sites with the inverted repeat
sequences TTGAT ATCAA. Fnr binding represses cyoABCDE
and cydAB operons and induces transcription from the operons
dmsABC (dimethyl sulfoxide/trimethylamine-N-oxide
reductase), frdABCD (fumarate reductase), and narGHJI
(nitrate reductase). Cross-talk between the two pathways at cyoABCDE
and cydAB is indicated by the broken arrow. Under microaerophilic
conditions as oxygen becomes limiting, cydAB is optimally active and
ArcA may successfully compete Fnr to activate the regulator under these
conditions (Bunn and Poyton,
1996).