A structural basis for mosaic protonic energy coupling.

to any given H + :P ratio. If the ratio is 3 and the permittivity 20, the distance is approximately 0.3nm. Unlike the electrochemical parameters, $“+ values do not depend on a cumulative ion effect; one proton alone at the right distance from a fixed charge will be at the potential required for synthetic activity. On these terms the rate of synthesis is determined by the number of charge<harge interactions effected in unit time, and respiratory control will be exerted through the turnover rate of the fixed-charge cycle (Archbold et al., 1979, 19806; Malpress, 19816) which provides the dynamic link between $ H + and its conversion into conformational change and intramembranal proton path activation. The coulombic hypothesis is a ‘local energized proton’ interpretation of mitochondrial energy transduction, and is therefore closer to the views of Williams (1 978) than to those of Mitchell (1979). But in many respects it may be radically differentiated from Williams’ prototype model, notably in its acceptance of the evidence for transmembrane proton pump activity as opposed to intermembrane proton release, in its explicit involvement of newly generated negative fixed-charges and of the diffuse double layer, by the electrostatic mechanism which it sees as the first step in the utilization of the energized protons in contrast to the kinetic ‘hydrating-dehydrating’ process of the prototype view (Fig. 3), and in its interpretation of Ca2+ movements.