The Bhopalator 137“6x9” b2861 The Cell Language Theory: Connecting Mind and MatterOne clear evidence for the existence of the t- and i-protons in mito-
chondria is provided by the two proton pathways or channels known to
operate in the cytochrome c oxidase (D- and K-pathways in Figures 3.39
and 3.40(a) and (b)). For example, the proton entering the oxygen reduc-
tion center of a cytochrome c oxidase from the matrix space (to be
denoted as H+N, where N indicates “negative”) via the D-pathway may
cross the membrane via the PLS (proton loading site) and ends up in the
inter-membrane space (i.e., as H+P, where P indicates “positive”), or may
be diverted to the heme a 3 site to participate in the oxygen reduction reac-
tion becoming a part of the water molecule formed (see Figure 3.40(a)).
In the former case, H+N exits the membrane to become H+P, which hence
would be an example of a t-H+. In the latter case, H+N enters the membrane
through the D-channel and combines with oxygen to form a water mole-
cule at the interior of cytochrome c oxidase, which would make it an
example of i-H+. So here is a clear case of a given proton acting as a t-H+
or as an i-H+, depending most likely on the conformational state of the
membrane enzyme system.
As pointed out in Figure 3.34, one major difference between the che-
miosmotic and conformon models of oxphos is that the chemiosmotic
model involves only one kind of protons (i.e., t-H+) that are directly linked
to both redox reaction and phosphorylation reaction, i.e., produced by
redox reaction and expelled into the intermembrane space later to be con-
sumed by phosphorylation reaction (with the H+/e- stoichiometry fixed to
1), whereas the conformon model postulates the existence of two kinds of
protons (i.e., i-H+ and t-H+), one participating in coupling two or more
intra-membrane enzyme systems moving within the membrane phase
(e.g., Complex IV and F 1 F 0 –ATP synthase) and the other traversing the
membrane and entering the inter-membrane space to create a trans-mem-
brane proton gradient. It was suggested above that the former kind of
protons be referred to as the Williams protons and the latter as Mitchell
proton. The Williams protons and Mitchell protons are closely related to
or identical with what some investigators call the chemical and physical
protons, respectively.
The structural similarity between the intracristae space of mitochon-
dria and the lumen of the thylakoid of chloroplast is shown in Table 3.15.
It is striking to notice that the intermembrane distances in the twob2861_Ch-03.indd 137 17-10-2017 11:46:51 AM