276 MHR • Unit 3 Molecular Genetics
more likely to bind to the operator and block
transcription. The tryptophan molecule thus
acts as a co-repressorto increase the affinity of
the repressor for the operator.
A prokaryote is a single-celled organism with a
relatively small genome and a short life span. Almost
all of its hereditary material is likely to be expressed
at some point during its life cycle. Therefore, gene
regulation in prokaryotes relies primarily on
mechanisms that can selectively turn off genes that
are not required under particular circumstances. In
contrast, eukaryotes may be multi-cellular organisms
with many specialized tissues. Most of the hereditary
material contained in any given eukaryotic cell is
therefore not expressed during the life of that cell.
As a result, eukaryotic cells are more dependent on
mechanisms that keep gene expression turned off
most of the time, and that turn on selected genes
only as they are needed.
Gene Expression in Eukaryotes
Figure 8.32 on the next page summarizes the
main steps in the protein synthesis pathway in
eukaryotes. These steps or control points (listed
below) offer more opportunities for control than
those available to prokaryotes.
pre-transcriptional control The cell controls the
extent to which DNA is exposed to transcription
enzymes, thus regulating the DNA’s availability
for transcription.
transcriptional control The cell controls whether
or not exposed DNA is transcribed into pre-mRNA.
post-transcriptional control The cell controls
the rate of processing of pre-mRNA into finished
mRNA.
translational control The cell manufactures
the mRNA, but then controls its transport to
ribosomes in the cytoplasm.
post-translational control The cell manufactures
the polypeptide product, but then modifies it
chemically or varies the rate at which it becomes
a functional protein. Another possibility is that
the cell may break the polypeptide product down
before it becomes a functional protein.
Like a prokaryotic cell, a eukaryotic cell can
save energy if regulation of gene expression occurs
sooner rather than later in the sequence of events
leading to protein synthesis. Nevertheless, a
eukaryotic cell is more likely to combine regulation
at the pre-transcriptional or transcriptional steps
with regulation at later steps. One reason for this
approach lies with the greater stability of mRNA
in eukaryotes. In prokaryotes, mRNA is typically
degraded within a few minutes after it is synthesized.
This is necessary because a gene that is expressed
will continue to be transcribed until it is turned off
by a regulatory mechanism. In contrast, mRNA in
eukaryotes tends to remain intact for hours or even
days, and it can be stored in the cell to be translated
at a later point. This means that a eukaryotic cell
can manufacture mRNA and then prevent it from
being translated until its protein product is required.
The following pages cover eukaryotic cell regulatory
mechanisms through the protein synthesis pathway.
Figure 8.31Co-repression in the E. coli trypoperon
tryp operon
repressor protein
binds to operator
P O
RNA polymerase
promoter and
operator sequences
tryp R gene
codes for
repressor
protein
tryp R
tryp D tryp B
tryp E tryp C tryp A
P O
tryp D tryp B
tryp E tryp C tryp A
repressor unbound
to operator
tryptophan co-repressor
binds to repressor
When tryptophan levels are low, the
repressor protein is present but does
not usually bind to the operator. RNA
polymerase is thus able to bind to the
promoter, so transcription takes place.
A
When tryptophan levels are high, tryptophan
binds to the repressor protein. As a result,
the repressor binds more readily to the
operator. When the repressor is bound to
the operator site, RNA polymerase cannot
bind and transcription will not proceed.