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Figure 17.1
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Concept 17.1: Genes specify proteins via
transcription and translation
RESULTS The wild-type strain required only the minimal medium for growth. The three classes of mutants had
different growth requirements
MM +
Citrulline
MM +
Arginine
(control)
Figure 17.2
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
CONCLUSION From the growth patterns of the mutants, Beadle and Tatum deduced that each mutant was unable
to carry out one step in the pathway for synthesizing arginine, presumably because it lacked the
necessary enzyme. Because each of their mutants was mutated in a single gene, they concluded that
each mutated gene must normally dictate the production of one enzyme. Their results supported the
one gene–one enzyme hypothesis and also confirmed the arginine pathway.
(Notice that a mutant can grow only if supplied with a compound made after the defective step.)
Enzyme
Gene A A A A A
Enzyme
Gene B B B B B
Citrulline Citrulline Citrulline Citrulline
Enzyme
Gene C C C C C
Arginine Arginine Arginine Arginine
• Translation
– Is the actual synthesis of a polypeptide, which
occurs under the direction of mRNA
– Occurs on ribosomes
DNA
TRANSCRIPTION
mRNA
Ribosome
TRANSLATION
Polypeptide
Figure 17.3a
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• In eukaryotes
– RNA transcripts are modified before becoming
true mRNA
Nuclear
envelope
TRANSCRIPTION DNA
Pre-mRNA
RNA PROCESSING
mRNA
Ribosome
TRANSLATION
(b) Eukaryotic cell. The nucleus provides a separate
Polypeptide compartment for transcription. The original RNA
transcript, called pre-mRNA, is processed in various
ways before leaving the nucleus as mRNA.
Figure 17.3b
DNA Gene 2
molecule
Gene 1
Gene 3
DNA strand 3′ 5′
(template) A C C A A A C C G A G T
TRANSCRIPTION
U G G U U U G G C U C A
mRNA 5′ 3′
Codon
TRANSLATION
Figure 17.6
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Concept 17.2: Transcription is the DNA-
directed synthesis of RNA: a closer look
Start point
Initiation. After RNA polymerase binds to
RNA polymerase 1
– Elongation the promoter, the DNA strands unwind, and
the polymerase initiates RNA synthesis at the
start point on the template strand.
5′ 3′
– Termination Unwound
3′
RNA DNA
Template strand of
5′
DNA transcript
2 Elongation. The polymerase moves downstream, unwinding the
DNA and elongating the RNA transcript 5′ → 3 ′. In the wake of
Rewound transcription, the DNA strands re-form a double helix.
RNA
5′ 3′
3′
3′ 5′
5′
RNA
transcript
3 Termination. Eventually, the RNA
transcript is released, and the
polymerase detaches from the DNA.
5′ 3′
3′ 5′
5′ 3′
Completed RNA
Figure 17.7 transcript
RNA
polymerase
T C C A A
A T
3′ C T U
3′ end
T
G
A U
G
G
A
C A E G C A C
5′ A
T A
A G G T T
Direction of transcription
5′ Template
(“downstream”)
strand of DNA
Newly made
RNA
• Transcription factors
– Help eukaryotic RNA polymerase recognize
promoter sequences TRANSCRIPTION DNA 1 Eukaryotic promoters
RNA PROCESSING Pre-mRNA
mRNA
TRANSLATION Ribosome
Polypeptide
Promoter
5′ T A T A A AA 3′
AT A T T T T
3′ 5′
TATA box Start point Template
DNA strand
2 Several transcription
factors
Transcription
factors
5′ 3′
3′ 5′
3 Additional transcription
factors
RNA polymerase II
Transcription factors
5′ 3′
3′ 5′ 5′
RNA transcript
Figure 17.8 Transcription initiation complex
Figure 17.9
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Split Genes and RNA Splicing
• RNA splicing
– Removes introns and joins exons
Polypeptide
mRNA 5′ Cap Poly-A tail
1 146
3′ UTR 3′ UTR
Figure 17.10
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Is carried out by spliceosomes in some cases
RNA transcript (pre-mRNA)
5′
Exon 1 Intron Exon 2
Protein
1
snRNA Other proteins
snRNPs
Spliceosome
2 5′
Spliceosome
components
Cut-out
intron
3
mRNA
5′
Figure 17.11 Exon 1 Exon 2
• In many cases
– Different exons code for the different domains in a
protein Gene
DNA
Exon 1 Intron Exon 2 Intron Exon 3
Transcription
RNA processing
Translation
Domain 3
Domain 2
Domain 1
mRNA
Ribosome
TRANSLATION
Polypeptide
Amino
Polypeptide acids
tRNA with
amino acid
Ribosome attached
Trp
Phe Gly
tRNA
C
C
GC G
A
Anticodon
A A A
U G G U U U G G C
5′ Codons 3′
mRNA
Figure 17.13
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Molecules of tRNA are not all identical
– Each carries a specific amino acid on one end
3′
– Is roughly L-shaped Amino acid A
C
attachment site C
A 5′
C G
G C
C G
U G
U A
A U
U C A U
* C A C AG U A G *
G A* C U C
*
C G U G U * C G A G G
* * U C * A G G
* G AG C
(a) Two-dimensional structure. The four base-paired regions and G C Hydrogen
three loops are characteristic of all tRNAs, as is the base sequence U A bonds
of the amino acid attachment site at the 3′ end. The anticodon triplet * G
A
is unique to each tRNA type. (The asterisks mark bases that have A* C
been chemically modified, a characteristic of tRNA.) * U
A G
A
Hydrogen
bonds
A AG
3′ 5′
Anticodon
Anticodon
(c) Symbol used
(b) Three-dimensional structure in this book
Figure 17.14b
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• A specific enzyme called an aminoacyl-tRNA
synthetase
– Joins each amino acid to the correct tRNA
Amino acid Aminoacyl-tRNA
synthetase (enzyme)
P P P Adenosine
1 Active site binds the
amino acid and ATP.
ATP
Pyrophosphate P Pi
Pi
Pi
Phosphates
tRNA
3 Appropriate
tRNA covalently
Bonds to amino
Acid, displacing P Adenosine
AMP. AMP
4 Activated amino acid
is released by the enzyme.
Aminoacyl tRNA
(an “activated
Figure 17.15 amino acid”)
mRNA
Ribosome
TRANSLATION
Polypeptide
Exit tunnel
Growing
polypeptide
tRNA
molecules
Large
subunit
E
P A
Small
subunit
5′
mRNA 3′
mRNA
binding site Small
subunit
tRNA
mRNA 3′
Codons
5′
(c) Schematic model with mRNA and tRNA. A tRNA fits into a binding site when its anticodon
base-pairs with an mRNA codon. The P site holds the tRNA attached to the growing
polypeptide. The A site holds the tRNA carrying the next amino acid to be added to the
polypeptide chain. Discharged tRNA leaves via the E site.
Figure 17.16c
– Elongation
– Termination
Initiator tRNA
GTP GDP
E A
mRNA
5′ 3′ 5′ 3′
Start codon
E E
P A P A
5′
3′ 3′
3′
5′ 5′
Stop codon
(UAG, UAA, or UGA)
1 When a ribosome reaches a stop 2 The release factor hydrolyzes 3 The two ribosomal subunits
codon on mRNA, the A site of the the bond between the tRNA in and the other components of
ribosome accepts a protein called the P site and the last amino the assembly dissociate.
a release factor instead of tRNA. acid of the polypeptide chain.
The polypeptide is thus freed
from the ribosome.
Figure 17.19
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Polyribosomes
• A number of ribosomes can translate a single
mRNA molecule simultaneously
– Forming a polyribosome Completed
Growing polypeptide
polypeptides
Incoming
ribosomal
subunits
Start of Polyribosom
eEnd of
mRNA mRNA
(5′ end) (3′ end)
(a) An mRNA molecule is generally translated simultaneously
by several ribosomes in clusters called polyribosomes.
Ribosomes
mRNA
0.1 µm
(b) This micrograph shows a large polyribosome in a prokaryotic
Figure 17.20a, b cell (TEM).
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Completing and Targeting the Functional Protein
• Polypeptide chains
– Undergo modifications after the translation
process
Ribosome
mRNA
Signal
peptide ER
membrane
Signal
Signal-
peptide
recognition Protein
removed
particle
(SRP) SRP
receptor
CYTOSOL protein
Translocation
ERLUMEN
complex
Figure 17.21
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Concept 17.5: RNA plays multiple roles in the
cell: a review
• RNA
– Can hydrogen-bond to other nucleic acid
molecules
– Can assume a specific three-dimensional
shape
– Has functional groups that allow it to act as a
catalyst
Table 17.1
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Concept 17.6: Comparing gene expression in
prokaryotes and eukaryotes reveals key differences
• Prokaryotic cells lack a nuclear envelope
DNA
mRNA
Polyribosome
Direction of 0.25 µm
RNA
transcription
polymerase
DNA
Polyribosome
Polypeptide
(amino end)
Ribosome
mRNA (5′ end)
Figure 17.22
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• In a eukaryotic cell
– The nuclear envelope separates transcription
from translation
– Extensive RNA processing occurs in the
nucleus
• Point mutations
– Are changes in just one base pair of a gene
mRNA mRNA
The mutant mRNA has
G A A G U A a U instead of an A in
one codon.
5′ 3′ 5′ 3′
mRNA
A U G A A G U U U GG C U A A
5′ 3′
Protein Met Lys Phe Gly Stop
Amino end
Carboxyl end
Base-pair substitution
No effect on amino acid sequence
U instead of C
A U G A A G U U U G G U U A A
A U G A A G U U U A G U U A A
A U G U U U G G C U A A
Figure 17.25 Met Phe Gly
Stop
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Mutagens
• Spontaneous mutations
– Can occur during DNA replication,
recombination, or repair
5′ RNA RNA
transcript polymerase
RNA PROCESSING Exon
2 In eukaryotes, the
RNA transcript
RNA transcript (pre-
(pre-mRNA)
mRNA) is spliced and
Intron
modified to produce
mRNA, which moves Aminoacyl-tRNA
p synthetase
from the nucleus to the Ca
cytoplasm. NUCLEUS
Amino
FORMATION OF
acid
INITIATION COMPLEX AMINO ACID ACTIVATION
CYTOPLASM 3 After leaving the tRNA
4 Each amino acid
nucleus, mRNA attaches attaches to its proper tRNA
to the ribosome. with the help of a specific
enzyme and ATP.
mRNA Growing
polypeptide
A Activated
ly-
Po amino acid
A
ly-
Ribosomal Po
subunits
p
Ca
5′
TRANSLATION
C 5 A succession of tRNAs
AC U
E A AC add their amino acids to
the polypeptide chain
AAA Anticodon
as the mRNA is moved
UG GUU U A U G through the ribosome
one codon at a time.
Codon (When completed, the
polypeptide is released
Ribosome
Figure 17.26 from the ribosome.)