The inner gate formed by Glu121, Glu122, His173, and Arg307 prevents water influx in the initial dark-adapted state. After the retinal isomerization reaction, the retinal twists and shifts toward Cys167 and induces an outward shift of TM3, originating at the kink introduced by Pro168. Isomerization of the retinal also induces a downward shift of the middle portion of TM7, especially at the retinal-attached residue, Lys296 (transition between the P1 and P2 intermediates). These movements are likely to induce water influx, by destabilizing the intracellular constriction, and further induce rearrangement of the surrounding helices (Open model).