Model for proposed ground-state conformation of lac permease, which
binds substrate with high affinity. For clarity, 6 of the 12 helices in
the permease are shown. Glu-126 (helix IV) and Arg-144 (helix V) are
charge-paired and with Cys-148 (helix V) comprise the major components
of the substrate binding site. Arg-302 (helix IX) is charge-paired with
Glu-325 (helix X), while protonated His-322 (helix X) interacts with
Glu-269 (helix VIII). Thus, the relevant H+ is shared by
His-322 and Glu-269. Also shown are the charge pair between Asp-240
(helix VII) and Lys-319 (helix X), which are not essential for the
mechanism. See text for further details regarding the transport
mechanism.