In complex III (cytochrome "bc1" complex or CoQH2-cytochrome "c" reductase; ), the Q-cycle contributes to the proton gradient by an asymmetric absorption/release of protons. Two electrons are removed from QH2 at the QO site and sequentially transferred to two molecules of cytochrome "c", a water-soluble electron carrier located within the intermembrane space. The two other electrons sequentially pass across the protein to the Qi site where the quinone part of ubiquinone is reduced to quinol. A proton gradient is formed by one quinol (<chem>2H+2e-</chem>) oxidations at the Qo site to form one quinone (<chem>2H+2e-</chem>) at the Qi site. (In total, four protons are translocated: two protons reduce quinone to quinol and two protons are released from two ubiquinol molecules.)