The Main Conclusions
* His291 is a probable pump element in CcO 
* Kinetic gating mechanism: ET → Fast PT (pumping H+) → Slow PT (chemical H+) 
* Proton pumping path involves: Glu242 → PRDa3/Arg438 → His291 
* 1e– /1H+ pumping stoichiometry
* Lys171B /Asp173B is a likely proton exit site 
* Absence of proton leaking is due to energy profile in exit proton channel 
 
A stepwise electron transfer from CuA to binuclear center is coupled to proton translocation from Glu242 to His291. [Popovic, Quenneville, Stuchebrukhov, JPC B, 109, 3616 (2005); BBA 1757, 1035, (2006)]
 
The schematics of the pumping model