130 The Cell Language Theory: Connecting Mind and Matterb2861 The Cell Language Theory: Connecting Mind and Matter “6x9”(3) It is now generally accepted that, when ATP synthase catalyzes the
formation of ATP from ADP and Pi, driven by the proton gradient
(regardless of whether the proton gradient is transmembrane as
assumed by the chemiosmotic model as depicted in Figure 3.32 or
intramembrane as assumed by the conformon model as depicted in
step 4 of Figure 3.29), the potential energy of the proton gradient is
first converted into the mechanical energy stored in F 0 [151, 161, 177,
211], which is then transferred to the g-subunit causing it to rotate,
leading to the rotary conformational wave propagation in F 1 resulting
in ATP synthesis followed by its expulsion from F 1 (see the figure on
the upper right-hand corner of Figure 3.35). Thus, the sequence of
events involved in the proton gradient-driven synthesis of ATP can
now be described as follows as proposed in [8, 210]:1
(Proton gradient) ↔ (Mechanical energy of the g- and e-subunits)
↔ 2
(Chemical energy of ATP) (3.39)Reaction (3.39) is known to be reversible (as indicated by the double
arrows) so that protons can be pumped across the mitochondrial inner
membrane (producing osmotic energy) using the chemical energy of ATP
hydrolysis. On the phenomenological level, therefore, the concept of che-
miosmotic coupling proposed by Mitchell may appear validated since
chemical and osmotic energies are indeed interconvertible. But this way of
looking at the problem is superficial. The heart of the problem concerns
not so much whether or not the process of chemiosmosis can occur in
mitochondria (which was known to occur in the organelle long before
1960 when the Mitchell hypothesis was formulated) but exactly how such
a process can occur on the molecular level. In other words, we must dis-
tinguish between the phenomenon of chemiosmosis and the molecular
mechanisms underlying chemiosmosis. On the phenomenological level,
the Mitchell hypothesis cannot be faulted. But it is on the level of molecu-
lar mechanisms underlying chemiosmosis that the Mitchell hypothesis
fails, as I have been pointing out over the last three decades [14, pp. 34–35;
7, pp. 60–61; 25, pp. 383–388] (see also [151, 161, 177, 209]).b2861_Ch-03.indd 130 17-10-2017 11:46:49 AM