tenth edition global edition - napa valley college … lecture...tenth edition global edition...
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BIOLOGY
Campbell • Reece • Urry • Cain • Wasserman • Minorsky • Jackson
© 2015 Pearson Education Ltd
TENTH EDITION
Global Edition
Lecture Presentation by
Nicole Tunbridge and
Kathleen Fitzpatrick
16Nucleic Acids and
Inheritance
© 2015 Pearson Education Ltd
Life’s Operating Instructions
a) In 1953, James Watson and Francis Crick introduced
an elegant double-helical model for the structure of
deoxyribonucleic acid, or DNA
b) Hereditary information is encoded in DNA and
reproduced in all cells of the body
c) This DNA program directs the development of
biochemical, anatomical, physiological, and
(to some extent) behavioral traits
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a) DNA is copied during DNA replication, and cells can
repair their DNA
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Concept 16.1: DNA is the genetic material
a) Early in the 20th century, the identification of the
molecules of inheritance loomed as a major challenge
to biologists
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The Search for the Genetic Material: Scientific Inquiry
a) When T. H. Morgan’s group showed that genes are
located on chromosomes, the two components of
chromosomes—DNA and protein—became
candidates for the genetic material
b) The role of DNA in heredity was first discovered
by studying bacteria and the viruses that
infect them
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Evidence That DNA Can Transform Bacteria
a) The discovery of the genetic role of DNA began with
research by Frederick Griffith in 1928
b) Griffith worked with two strains of a bacterium, one
pathogenic and one harmless
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a) When he mixed heat-killed remains of the pathogenic
strain with living cells of the harmless strain, some
living cells became pathogenic
b) He called this phenomenon transformation, now
defined as a change in genotype and phenotype due
to assimilation of foreign DNA
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Figure 16.2
Living S cells(pathogeniccontrol)
Experiment
Results
Living R cells(nonpathogeniccontrol)
Heat-killed S cells(nonpathogeniccontrol)
Mouse dies Mouse healthy Mouse healthy Mouse dies
Mixture of heat-killed S cells andliving R cells
Living S cells
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a) In 1944, Oswald Avery, Maclyn McCarty, and Colin
MacLeod announced that the transforming substance
was DNA
b) Many biologists remained skeptical, mainly because
little was known about DNA
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Evidence That Viral DNA Can Program Cells
a) More evidence for DNA as the genetic material came
from studies of viruses that infect bacteria
b) Such viruses, called bacteriophages (or phages),
are widely used in molecular genetics research
c) A virus is DNA (sometimes RNA) enclosed by a
protective coat, often simply protein
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Figure 16.3
Phage
head
DNA
Tail
sheath
Tail fiber
Genetic
material
Bacterial
cell
100 n
m
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a) In 1952, Alfred Hershey and Martha Chase showed
that DNA is the genetic material of a phage known as
T2
b) They designed an experiment showing that only one
of the two components of T2 (DNA or protein) enters
an E. coli cell during infection
c) They concluded that the injected DNA of the phage
provides the genetic information
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Figure 16.4
Experiment
Batch 1: Radioactive sulfur (35S) in phage protein
Batch 2: Radioactive phosphorus (32P) in phage DNA
Labeled phagesinfect cells.
Agitation frees outsidephage parts from cells.
Centrifuged cellsform a pellet.
Radioactivity(phage protein)found in liquid
Pellet
Centrifuge
Radioactiveprotein
RadioactiveDNA
Radioactivity (phageDNA) found in pellet
Centrifuge
Pellet
1 2
4
3
4
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Figure 16.4a
Experiment
Batch 1: Radioactive sulfur (35S) in phage protein
Labeled phagesinfect cells.
Agitation frees outside phage parts from cells.
Centrifuged cellsform a pellet.
Radioactivity(phage protein)found in liquid
Radioactiveprotein
Pellet
Centrifuge
1 2 3
4
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Figure 16.4b
Pellet
Centrifuge
Batch 2: Radioactive phosphorus (32P) in phage DNA
RadioactiveDNA
Radioactivity (phage DNA) found in pellet
Labeled phagesinfect cells.
Agitation frees outside phage parts from cells.
Centrifuged cellsform a pellet.
Experiment
1 2 3
4
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Additional Evidence That DNA Is the Genetic Material
a) It was known that DNA is a polymer of nucleotides,
each consisting of a nitrogenous base, a sugar, and a
phosphate group
b) In 1950, Erwin Chargaff reported that DNA
composition varies from one species to the next
c) This evidence of diversity made DNA a more credible
candidate for the genetic material
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Figure 16.5
5′ end
Thymine (T)
Adenine (A)
Cytosine (C)
Guanine (G)
Nitrogenous
bases
Sugar–
phosphate
backbone
3′ end
Nitrogenous
base
Sugar
(deoxyribose)DNA
nucleotide
Phosphate
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a) Two findings became known as Chargaff’s rules
a)The base composition of DNA varies between species
b)In any species the number of A and T bases are equal
and the number of G and C bases are equal
b) The basis for these rules was not understood until the
discovery of the double helix
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Building a Structural Model of DNA: Scientific Inquiry
a) After DNA was accepted as the genetic material, the
challenge was to determine how its structure
accounts for its role in heredity
b) Maurice Wilkins and Rosalind Franklin were using a
technique called X-ray crystallography to study
molecular structure
c) Franklin produced a picture of the DNA molecule
using this technique
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Figure 16.6
(a) Rosalind Franklin (b) Franklin’s X-ray diffraction
photograph of DNA
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a) Franklin’s X-ray crystallographic images of DNA
enabled Watson to deduce that DNA was helical
b) The X-ray images also enabled Watson to deduce the
width of the helix and the spacing of the nitrogenous
bases
c) The pattern in the photo suggested that the DNA
molecule was made up of two strands, forming a
double helix
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Figure 16.7
(a) Key features of
DNA structure
(b) Partial chemical structure
0.34 nm
3′ end
5′ endT
T
T
A
A
A
C
C
C
G
G
G
AT1 nm
TA
C G
CG
AT3.4 nm
CG
CG
C G
C G
3′ end
5′ end
Hydrogen bond
T A
G C
A T
C G
(c) Space-filling
model
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Figure 16.7a
(a) Key features of DNA structure
0.34 nm
1 nm
3.4 nm
T
T
T
A
A
A
C
C
C
G
G
G
AT
TA
C G
CG
AT
CG
CG
C G
C G
© 2015 Pearson Education Ltd
Figure 16.7b
(b) Partial chemical structure
3′ end
5′ end
3′ end
5′ end
Hydrogen bond
T A
G C
A T
C G
© 2015 Pearson Education Ltd
a) Watson and Crick built models of a double helix to
conform to the X-rays and chemistry of DNA
b) Franklin had concluded that there were two outer
sugar-phosphate backbones, with the nitrogenous
bases paired in the molecule’s interior
c) Watson built a model in which the backbones were
antiparallel (their subunits run in opposite directions)
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a) At first, Watson and Crick thought the bases paired
like with like (A with A, and so on), but such pairings
did not result in a uniform width
b) Instead, pairing a purine with a pyrimidine resulted in
a uniform width consistent with the X-ray data
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Figure 16.UN02
Purine + purine: too wide
Pyrimidine + pyrimidine: too narrow
Purine + pyrimidine: width
consistent with X-ray data
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a) Watson and Crick reasoned that the pairing was more
specific, dictated by the base structures
b) They determined that adenine (A) paired only with
thymine (T), and guanine (G) paired only with
cytosine (C)
c) The Watson-Crick model explains Chargaff’s rules: in
any organism the amount of A = T, and the amount of
G = C
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Figure 16.8
Sugar
Sugar
Adenine (A)
Sugar
Sugar
Thymine (T)
Cytosine (C)Guanine (G)
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Concept 16.2: Many proteins work together in DNA replication and repair
a) The relationship between structure and function is
manifest in the double helix
b) Watson and Crick noted that the specific base pairing
suggested a possible copying mechanism for genetic
material
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The Basic Principle: Base Pairing to a Template Strand
a) Since the two strands of DNA are complementary,
each strand acts as a template for building a new
strand in replication
b) In DNA replication, the parent molecule unwinds, and
two new daughter strands are built based on base-
pairing rules
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Figure 16.9-2
(a) Parental
molecule(b) Separation of parental
strands into templates
A
A
A
T
T
T
G
G C
C
A
A
A
T
T
T
G
G C
C
© 2015 Pearson Education Ltd
Figure 16.9-3
(a) Parental
molecule(b) Separation of parental
strands into templates
A
A
A
T
T
T
G
G C
C
A
A
A
T
T
T
G
G C
C
A
A
A
T
T
T
G
G C
C
A
A
A
T
T
T
G
G C
C
(c) Formation of new strands
complementary to template
strands
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a) Watson and Crick’s semiconservative model of
replication predicts that when a double helix
replicates, each daughter molecule will have one old
strand (derived or “conserved” from the parent
molecule) and one newly made strand
b) Competing models were the conservative model (the
two parent strands rejoin) and the dispersive model
(each strand is a mix of old and new)
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Figure 16.10
(a) Conservative
model
(b) Semiconserva-
tive model
(c) Dispersive
model
Parent cell
First
replication
Second
replication
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a) Experiments by Matthew Meselson and Franklin Stahl
supported the semiconservative model
b) They labeled the nucleotides of the old strands with a
heavy isotope of nitrogen, while any new nucleotides
were labeled with a lighter isotope
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a) The first replication produced a band of hybrid DNA,
eliminating the conservative model
b) A second replication produced both light and hybrid
DNA, eliminating the dispersive model and supporting
the semiconservative model
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Figure 16.11
Bacteria cultured
in medium with 15N
(heavy isotope)
Experiment
Results
Conclusion
Bacteria transferred
to medium with 14N
(lighter isotope)
DNA sample
centrifuged
after first
replication
DNA sample
centrifuged
after second
replication
Less dense
More dense
Predictions: First replication Second replication
Conservative
model
Semiconservative
model
Dispersive
model
1 2
43
© 2015 Pearson Education Ltd
Figure 16.11a
Bacteria cultured
in medium with 15N
(heavy isotope)
Experiment
Results
Bacteria transferred
to medium with 14N
(lighter isotope)
DNA samplecentrifugedafter firstreplication
DNA samplecentrifugedafter secondreplication
Less dense
More dense
1 2
3 4
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Figure 16.11b
Conclusion
Predictions: First replication Second replication
Conservative
model
Semiconservative
model
Dispersive
model
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DNA Replication: A Closer Look
a) The copying of DNA is remarkable in its speed and
accuracy
b) More than a dozen enzymes and other proteins
participate in DNA replication
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Getting Started
a) Replication begins at particular sites called origins of
replication, where the two DNA strands are
separated, opening up a replication “bubble”
b) A eukaryotic chromosome may have hundreds or
even thousands of origins of replication
c) Replication proceeds in both directions from each
origin, until the entire molecule is copied
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Figure 16.12
Origin of
replication
0.5
µm
0.2
5 µ
m
Bacterial
chromosome
Two daughter
DNA molecules
Replication
bubble
Parental (template)
strandDaughter
(new) strand
Replication
fork
Double-
stranded
DNA molecule
(a) Origin of replication in an E. coli cell (b) Origins of replication in a eukaryotic cell
Origin of
replication Eukaryotic chromosome
Double-stranded
DNA molecule
Parental (template)
strandDaughter (new)
strand
Replication
forkBubble
Two daughter DNA molecules
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Figure 16.12a
0.5
µm
Origin of
replication
Bacterial
chromosome
Two daughter
DNA molecules
Replication
bubble
Parental (template) strand
Daughter
(new) strand
Replication
forkDouble-stranded
DNA molecule
(a) Origin of replication in an E. coli cell
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Figure 16.12b
0.2
5 µ
m
(b) Origins of replication in a eukaryotic cell
Origin of replication Eukaryotic chromosome
Double-stranded
DNA molecule
Parental (template) strand
Daughter (new)
strand
Replication
forkBubble
Two daughter DNA molecules
© 2015 Pearson Education Ltd
a) At the end of each replication bubble is a replication
fork, a Y-shaped region where
new DNA strands are elongating
b)Helicases are enzymes that untwist the double helix
at the replication forks
c) Single-strand binding proteins bind to and stabilize
single-stranded DNA
d)Topoisomerase corrects “overwinding” ahead of
replication forks by breaking, swiveling, and rejoining
DNA strands
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Figure 16.13
Topoisomerase
Primase
RNA
primer
Replication
fork
5′
3′
5′
5′
3′
Helicase
Single-strand binding
proteins
3′
© 2015 Pearson Education Ltd
a) DNA polymerases cannot initiate synthesis of a
polynucleotide; they can only add nucleotides to an
existing 3′ end
b) The initial nucleotide strand is a short RNA primer
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a) An enzyme called primase can start an RNA chain
from scratch and adds RNA nucleotides one at a time
using the parental DNA as a template
b) The primer is short (5–10 nucleotides long), and the
3′ end serves as the starting point for the new DNA
strand
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Synthesizing a New DNA Strand
a) Enzymes called DNA polymerases catalyze the
elongation of new DNA at a replication fork
b) Most DNA polymerases require a primer and a DNA
template strand
c) The rate of elongation is about 500 nucleotides per
second in bacteria and 50 per second in human cells
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a) Each nucleotide that is added to a growing DNA
strand is a nucleoside triphosphate
b) dATP supplies adenine to DNA and is similar to the
ATP of energy metabolism
c) The difference is in their sugars: dATP has
deoxyribose while ATP has ribose
d) As each monomer of dATP joins the DNA strand, it
loses two phosphate groups as a molecule of
pyrophosphate
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Figure 16.14
5′ 5′3′
3′
5′ 5′
New strand Template strand
Sugar
Phosphate Base
OH
A T
C
C
G
G
A
C
Nucleotide
DNApoly-
merase
OH
Pyro-phosphate
2 P i
P iP
3′
3′
A T
C
C
G
G
A
C
T
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Antiparallel Elongation
a) The antiparallel structure of the double helix affects
replication
b) DNA polymerases add nucleotides only to the free 3′
end of a growing strand; therefore, a new DNA strand
can elongate only in the 5′ to 3′ direction
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Figure 16.15
Leading strand Lagging strand
Leading strand
Parental DNA
Lagging strand
Primer
Overview
Origin of replication
Overall
directions
of replication Origin of replication
RNA primer
Sliding clamp
DNA pol III
Continuous
elongation in the
5′ to 3′ direction
5′
5′
5′
5′
5′
5′
3′ 3′
3′3′
3′
3′
DNA pol III starts to
synthesize leading
strand.
1
2
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Figure 16.15a
Leading strand Lagging strand
Leading strandLagging strand
Primer
Overview
Origin of replication
Overall directions
of replication
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Figure 16.15b
Parental DNA
Origin of replication
RNA primer
Sliding clamp
DNA pol III
5′
5′
5′
3′ 3′
3′
DNA pol III starts to
synthesize leading
strand.
1
5′
3′
5′
3′
5′
3′
2 Continuous
elongation in the
5′ to 3′ direction
© 2015 Pearson Education Ltd
a) Along one template strand of DNA, the DNA
polymerase synthesizes a leading strand
continuously, moving toward the replication fork
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a) To elongate the other new strand, called the lagging
strand, DNA polymerase must work in the direction
away from the replication fork
b) The lagging strand is synthesized as a series of
segments called Okazaki fragments, which are
joined together by DNA ligase
© 2015 Pearson Education Ltd
Figure 16.16Overview
12
1
1
2
1
2
1
1
2
5′
3′
3′
5′5′3′
3′
3′ 3′5′
5′5′
5′3′
3′
5′3′
5′3′
5′3′
5′3′
5′3′
5′3′
1
2 5
6
4
3
Origin of replication
Lagging
strand
Leading
strand
Lagging
strand
Leading
strandOverall directions
of replication
RNA primer
for fragment 2
Okazaki
fragment 2DNA pol III
makes Okazaki
fragment 2.
DNA pol I
replaces RNA
with DNA.
DNA ligase
forms bonds
between DNA
fragments.
Overall direction of replication
Okazaki
fragment 1
DNA pol III
detaches.
RNA primer
for fragment 1
Template
strand
DNA pol III
makes Okazaki
fragment 1.
Origin of
replication
Primase makes
RNA primer.
5′
© 2015 Pearson Education Ltd
Figure 16.16a
12
Overview
Origin of replication
Lagging
strand
Leading
strand
Lagging
strand
Leading
strandOverall directions
of replication
© 2015 Pearson Education Ltd
Figure 16.16b-1
5′
3′
5′5′3′
3′Template
strand
Origin of
replication
Primase makes
RNA primer.
1
© 2015 Pearson Education Ltd
Figure 16.16b-2
5′
3′
5′5′3′
3′Template
strand
Origin of
replication
Primase makes
RNA primer.
1
5′3′
5′
3′
5′3′
RNA primer
for fragment 1
DNA pol III
makes Okazaki
fragment 1.
1
2
© 2015 Pearson Education Ltd
Figure 16.16b-3
5′
3′
5′5′3′
3′Template
strand
Origin of
replication
Primase makes
RNA primer.
1
5′3′
5′
3′
5′3′
RNA primer
for fragment 1
DNA pol III
makes Okazaki
fragment 1.
1
2
1
3
3′
3′5′
5′
Okazaki
fragment 1
DNA pol III
detaches.
© 2015 Pearson Education Ltd
DNA pol III
makes Okazaki
fragment 2.
Figure 16.16c-1
2
5′
3′
5′3′
4
RNA primer
for fragment 2Okazaki
fragment 2
1
© 2015 Pearson Education Ltd
Figure 16.16c-2
DNA pol III
makes Okazaki
fragment 2.2
5′
3′
5′3′
4
RNA primer
for fragment 2Okazaki
fragment 2
1
5′3′
DNA pol I
replaces RNA
with DNA.
5′3′
1
2
5
© 2015 Pearson Education Ltd
Figure 16.16c-3
DNA ligaseforms bondsbetween DNAfragments.
Overall direction of replication
DNA pol III
makes Okazaki
fragment 2.2
5′
3′
5′3′
4
RNA primer
for fragment 2Okazaki
fragment 2
1
5′3′
DNA pol I
replaces RNA
with DNA.
5′3′
1
2
5
6
5′3′
5′3′
1
2
© 2015 Pearson Education Ltd
Figure 16.17
Overview
5′
3′
Lagging
strand
Leading
strand
Leading
strand
Lagging
strandLeading strand
Leading strand
template
Origin of
replication
Overall directions
of replication
5′
3′
5′3′
5′
5′
3′
3′3′
Single-strand
binding proteins
Helicase
Parental
DNA
DNA pol III
Primer
Primase
Lagging
strand
Lagging strand
template
DNA pol III
DNA pol I5′
DNA ligase
123
4
5
© 2015 Pearson Education Ltd
Figure 16.17a
Overview
Lagging
strand
Leading
strand
Leading
strand
Lagging
strand
Origin of
replication
Overall directions
of replication
© 2015 Pearson Education Ltd
Figure 16.17b
5′
5′
3′
3′3′
Single-strand
binding proteins
Helicase
Parental
DNA
DNA pol III
Primer
Primase
Leading
strand
Leading strand
template
5
© 2015 Pearson Education Ltd
Figure 16.17c
4
3 2 1
5′
5′3′
5′
3′
Lagging
strand
DNA pol III
DNA pol I DNA ligase
Lagging strand
template
© 2015 Pearson Education Ltd
The DNA Replication Complex
a) The proteins that participate in DNA replication form a
large complex, a “DNA replication machine”
b) The DNA replication machine may be stationary
during the replication process
c) Recent studies support a model in which DNA
polymerase molecules “reel in” parental DNA and
“extrude” newly made daughter DNA molecules
© 2015 Pearson Education Ltd
Figure 16.18
Leading strand template
5′
5′
5′
5′3′
3′
3′
3′
3′ 3′
5′
5′
Leading strand
Lagging strand
Lagging
strand
template
DNA pol III
Connecting proteinHelicase
Parental DNA
DNA pol III
© 2015 Pearson Education Ltd
Proofreading and Repairing DNA
a) DNA polymerases proofread newly made DNA,
replacing any incorrect nucleotides
b) In mismatch repair of DNA, repair enzymes correct
errors in base pairing
c) DNA can be damaged by exposure to harmful
chemical or physical agents such as cigarette smoke
and X-rays; it can also undergo spontaneous
changes
d) In nucleotide excision repair, a nuclease cuts out
and replaces damaged stretches of DNA
© 2015 Pearson Education Ltd
Figure 16.19-2
Nuclease
5′
5′
5′
5′
3′
3′
3′
3′
5′
5′
3′
3′
DNA
polymerase
© 2015 Pearson Education Ltd
Figure 16.19-3
Nuclease
5′
5′
5′
5′
3′
3′
3′
3′
5′
5′
3′
3′
DNA
polymerase
DNA
ligase
5′
3′5′
3′
© 2015 Pearson Education Ltd
Evolutionary Significance of Altered DNA Nucleotides
a) Error rate after proofreading repair is low but
not zero
b) Sequence changes may become permanent and can
be passed on to the next generation
c) These changes (mutations) are the source of the
genetic variation upon which natural selection
operates
© 2015 Pearson Education Ltd
Replicating the Ends of DNA Molecules
a) Limitations of DNA polymerase create problems for
the linear DNA of eukaryotic chromosomes
b) The usual replication machinery provides no way to
complete the 5′ ends, so repeated rounds of
replication produce shorter DNA molecules with
uneven ends
c) This is not a problem for prokaryotes, most of which
have circular chromosomes
© 2015 Pearson Education Ltd
Figure 16.20
Ends of parental
DNA strands
Lagging strand
Parental strand
RNA primer
Last fragmentNext-to-last
fragment
Lagging strand
Leading strand
Removal of primers and
replacement with DNA
where a 3′ end is available
3′
5′
5′
Second round
of replication
Further rounds
of replication
New lagging strand
New leading strand
Shorter and shorter daughter molecules
5′
3′
3′
5′
3′
5′
3′
© 2015 Pearson Education Ltd
Figure 16.20a
Ends of parental
DNA strands
Lagging strand
Parental strand
RNA primer
Last fragment
Next-to-last
fragment
Lagging strand
Leading strand
Removal of primers and
replacement with DNA
where a 3′ end is available
3′
5′
5′
3′
5′
3′
© 2015 Pearson Education Ltd
Figure 16.20b
5′
Second round
of replication
Further rounds
of replication
New lagging strand
New leading strand
Shorter and shorter daughter molecules
5′
3′
3′
5′
3′
© 2015 Pearson Education Ltd
a) Eukaryotic chromosomal DNA molecules have
special nucleotide sequences at their ends called
telomeres
b) Telomeres do not prevent the shortening of DNA
molecules, but they do postpone the erosion of genes
near the ends of DNA molecules
c) It has been proposed that the shortening of telomeres
is connected to aging
© 2015 Pearson Education Ltd
a) If chromosomes of germ cells became shorter in
every cell cycle, essential genes would eventually be
missing from the gametes they produce
b) An enzyme called telomerase catalyzes the
lengthening of telomeres in germ cells
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a) The shortening of telomeres might protect cells from
cancerous growth by limiting the number of cell
divisions
b) There is evidence of telomerase activity in cancer
cells, which may allow cancer cells to persist
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Concept 16.3: A chromosome consists of a DNA molecule packed together with proteins
a) The bacterial chromosome is a double-stranded,
circular DNA molecule associated with a small
amount of protein
b) Eukaryotic chromosomes have linear DNA molecules
associated with a large amount
of protein
c) In a bacterium, the DNA is “supercoiled” and found in
a region of the cell called the nucleoid
© 2015 Pearson Education Ltd
a) In the eukaryotic cell, DNA is precisely combined with
proteins in a complex called chromatin
b) Chromosomes fit into the nucleus through an
elaborate, multilevel system of packing
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Figure 16.22
DNA, the
double helixHistones Nucleosomes,
or “beads on
a string”
(10-nm fiber)
30-nm fiberLooped
domains
(300-nm fiber)
Metaphase
chromosome
DNA
double helix
(2 nm in diameter)
Nucleosome
(10 nm in diameter)30-nm fiber
Loops Scaffold
HistonesHistone tail
H1
300-nm
fiber
Chromatid
(700 nm)
Replicated
chromosome
(1,400 nm)
© 2015 Pearson Education Ltd
Figure 16.22a
DNA, the
double helix
Histones Nucleosomes, or “beads on
a string” (10-nm fiber)
DNA
double helix
(2 nm in diameter)
Nucleosome
(10 nm in diameter)
HistonesHistone tail
H1
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Figure 16.22b
30-nm fiber
Looped domains
(300-nm fiber)
Metaphase
chromosome
30-nm fiber
Loops Scaffold
300-nm fiber
Chromatid
(700 nm)
Replicated
chromosome
(1,400 nm)
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a) Chromatin undergoes changes in packing during the
cell cycle
b) At interphase, some chromatin is organized into a 10-
nm fiber, but much is compacted into a 30-nm fiber,
through folding and looping
c) Interphase chromosomes occupy specific restricted
regions in the nucleus and the fibers of different
chromosomes do not become entangled
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a) Most chromatin is loosely packed in the nucleus
during interphase and condenses prior to mitosis
b) Loosely packed chromatin is called euchromatin
c) During interphase a few regions of chromatin
(centromeres and telomeres) are highly condensed
into heterochromatin
d) Dense packing of the heterochromatin makes it
difficult for the cell to express genetic information
coded in these regions
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a) Histones can undergo chemical modifications that
result in changes in chromatin organization
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Figure 16.UN01
Base Percentage
Source of DNA Adenine Guanine Cytosine Thymine
Sea urchin 32.8 17.7 17.3 32.1
Salmon 29.7 20.8 20.4 29.1
Wheat 28.1 21.8 22.7
E. coli 24.7 26.0
Human 30.4 30.1
Ox 29.0
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Figure 16.UN02
Purine + purine: too wide
Pyrimidine + pyrimidine: too narrow
Purine + pyrimidine: width
consistent with X-ray data
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Figure 16.UN03
Sugar-phosphate
backbone
Nitrogenous
bases
Hydrogen bond
G
G
G
G
G
C
C
C
C
C
A T
T A
A T
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Figure 16.UN04
DNA pol III synthesizes
leading strand continuously
Parental
DNA
Helicase
Primase synthesizes
a short RNA primer
3′
5′
DNA pol I replaces the RNA
primer with DNA nucleotides
3′
5′
3′
5′
DNA pol III starts DNA
synthesis at 3′ end of primer,
continues in 5′ → 3′ direction
Lagging strand synthesized
in short Okazaki fragments,
later joined by DNA ligase
Origin of
replication