1. cardiolipin which aggregates the respiratory system together with the ATP
synthase. It also restricts the H
+ coming from the respiratory complex close to
ATP synthase and facilitate the H
+ transfer to the synthase [144],
2. cytochromes that pump out H
+ much faster than the normal pace of nonalkaliphilic organisms [161, 175] and cytochromes like cytochrome c-550 with
high electron retention capacity [174, 177], and
3. ATP synthase which is efficient in translocating H
+ to its catalytic core (i.e.
inhibition of H
+ leakage) [95, 170, 173].
The rapid pumping of H
+ by the respiratory complex and the restriction of these
H
+ close to the surface of the aggregate create a microenvironment with high pmf
which promotes the synthesis of ATP (Fig. 10). The H
+ translocated by the ATP
synthase during ATP synthesis replenishes the H
+ pumped out by the respiratory
complex, which contributes to maintain the low cytoplasmic pH. This microcircuit in
the microenvironment produces ATP approximately seven times faster than that of
neutralophiles [160, 177], and this may be one of the reasons why alkaliphiles grow
faster and denser than neutralophiles.
Fig. 10 An illustration of the high pmf microenvironment created by cardiolipin-induced aggregation of the respiratory complexes (complex III and IV). Quinone oxidoreductase (QO, NDH-2)
transfers electrons to menaquinone (MQ) pool from which the electrons move to the menaquinol:
cytochrome c (Complex III) and cytochrome c oxidase (complex IV). The rapid pumping of H
+ by
the respiratory complexes and the retention of electrons by the cytochrome c-550 contribute to high
pmf with more negative charges on the membrane facing the cytoplasm than the outer membrane
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