Reaction scheme of translocation as discussed in
the text. EF-G (green) is depicted in three conformations: the
GTP-bound form, an intermediate GDP-bound form on the ribosome, and the
GDP-bound form that dissociates from the ribosome (step 6); the latter
conformation probably is represented in the crystal structures of
EF-G-GDP and nucleotide-free EF-G (11, 12). Pi is omitted
because it is not known when after GTP hydrolysis it is released. The
transition state of the ribosome, formed in step 3, is symbolized by an
altered conformation of the small ribosomal subunit; of course,
alterations of the large subunit or an altered arrangement of the
subunits relative to each other also may be involved. A, P (P*), and E
denote the tRNA binding sites on the two subunits and are indicated
when occupied.