Chapter 8 Protein Synthesis • MHR 265
Each active ribosome has a binding site for the
mRNA transcript and three binding sites for tRNA
molecules: the P site, which holds one aa-tRNA
and the growing chain of amino acids; the A site,
which holds the tRNA bringing the next amino
acid to be added to the chain; and the E site,
which releases the tRNA molecules back into
the cytoplasm. These binding sites are provided
by complementary nucleotide sequences on the
different RNA molecules.
Like the transcription process, translation can
be examined as a sequence of steps: initiation,
elongation, and termination. These three steps
are examined in more detail below.
Initiation
Translation is initiated when an mRNA molecule
reaches the cytoplasm of the cell. A sequence of
nucleotides at the 5 ′end of the mRNA molecule
binds to a portion of the rRNA strand in a small
ribosomal sub-unit. A special initiator tRNA
molecule also binds to the ribosomal-mRNA
complex. This molecule has the anticodon UAC (to
complement the start codon AUG) and carries the
amino acid methionine. The resulting combination
binds to a large ribosomal sub-unit in order to
complete the assembly of the translation initiation
complex as illustrated in Figure 8.18.
Figure 8.17The two sub-units in an active ribosome form
binding sites for the mRNA transcript and the aa-tRNA
molecules involved in protein synthesis. Ribosomal RNA
molecules are not specific to any polypeptide product,
which means that the same ribosome can be used to
synthesize any protein.
Figure 8.18At the end of the initiation stage, the two
ribosomal sub-units have come together to create an active
ribosome bound to a strand of mRNA. An initiator tRNA sits
at the P site. Both the A site and the E site are vacant.
As part of the initiation process, the ribosomal
sub-units must bind at precisely the right place
on the mRNA molecule. If the only signal for
translation initiation was the start codon AUG,
a tRNA anticodon could incorrectly bind to the
sequence AUG at any point along the mRNA
strand. Base pairing between the mRNA molecule
and the nucleotide sequence in rRNA help resolve
this problem.
To understand how this process works,
remember that each mRNA molecule has a leader
sequence ahead of the start codon. This leader
sequence interacts with the rRNA in the small
ribosomal sub-unit to help establish the correct
starting place for translation. This interaction takes
place before the large ribosomal sub-unit joins the
initiation complex, with the result that the initial
anticodon binding site is correctly established
before translation begins.
Elongation
Once the initiation step is complete, the synthesis
of a polypeptide involves repeating three general
steps in a cycle. First the mRNA codon exposed
in the A binding site forms a base pair with the
anticodon of an incoming aa-tRNA molecule. Then
enzymes and other molecules in the large sub-unit
catalyze the formation of a peptide bond that joins
the last amino acid in the growing peptide chain
to the new amino acid. At the same time, the
polypeptide chain is transferred from the tRNA
in the P site to the tRNA in the A site.
In the third step, the ribosome moves a distance
of three nucleotides along the mRNA molecule.
This shift, which results in the ribosome assembly
initiator tRNA
in P site
met
EA
5 ′
3 ′
large ribosomal
sub-unit
small ribosomal
sub-unit
uac
aug
P site
A site
large ribosomal
sub-unit
small ribosomal
sub-unit
mRNA
binding
site
E site
Many antibiotics work by paralyzing prokaryotic ribosomes.
The structural differences between prokaryotic and
eukaryotic ribosomes mean that these antibiotics can
kill bacteria without harming the patient.