88 The Cell Language Theory: Connecting Mind and Matterb2861 The Cell Language Theory: Connecting Mind and Matter “6x9”Figure 3.18 The postulated existence of the electron-, proton-, and phosphoron-transfer
complexes in the mitochondrial inner membrane. The mono-negatively charged phos-
phoryl group, i.e., PO 3 - , was named the phosphoron in 1976 [8]. PTC is also called the
transprotonase [144], in analogy to the well-known transhydrogenase. Adopted from [8].
ETC = electron-transfer complex; PTC = proton-transfer complex; OSA = oligomycin-
sensitive ATPase, equivalent to the F 0 F 1 –ATPase/synthase [242]. The combination of ETC,
PTC, and OSA depicted here can be viewed as an abstract version of the supermolecule
shown in Figure 3.4(a) or the oxphosome defined in Table 3.5.ETC PTC OSAε– H+ H+ φ–(see Figure 3.29, Step 4), will be referred to as the second-order pseudolink-
age, in analogy to the first-order and second-order kinetics in chemistry. It
is possible that in mitochondrial inner membrane, a third-order pseudolink-
age is common, e.g., among ETC, PTC, and OSA mediated by intramem-
brane protons of Williams (see Section 3.3.6) (as opposed to the Mitchell’s
protons in the extramembrane bulk phases) as schematically depicted in
Figure 3.18. In ETC, the electron and the proton are truly linked; in PTC,
there are postulated to be at least two truly linked proton binding sites (e.g.,
aspartate, glutamate); and, in OSC, protons and phosphorons are thought to
be truly linked in such a way that when the amount of protons in PTC is kept
constant, ETC and OSA become pseudolinked, so that respiration can drive
the synthesis of ATP. When the intramembrane protons (referred on top as
the Williams protons, see Section 3.3.6) equilibrate with bulk phase protons
due to lipid-soluble anions such as dinitrophenol, the amount of the intram-
embrane protons becomes variable (depending on the extramembrane con-
centrations) while their chemical activity remains constant, thereby breaking
down the pseudolinkage and resulting in uncoupling.
The best known example of the true linkage is the Bohr effect in
hemoglobin [168] as described in Figure 3.16, where O 2 and H+ are theb2861_Ch-03.indd 88 17-10-2017 11:46:24 AM