CH. 12 CELL DIVISION
Life is based on the reproduction of
cells, or cell division
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FIGURE 12.1
In unicellular organisms, division of one
cell reproduces the entire organism
Multicellular organisms depend on cell
division for
 Development from a fertilized cell
 Growth
 Repair
© 2011 Pearson Education, Inc.
FIGURE 12.2
100 m
(a) Reproduction
200 m
(b) Growth and
development
20 m
(c) Tissue renewal
DAUGHTER CELLS
Most cell division results in daughter
cells with identical genetic
information, DNA
The exception is meiosis, a special
type of division that can produce
sperm and egg cells
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GENETIC MATERIAL
All the DNA in a cell constitutes the cell’s
genome
A genome can consist of a single DNA
molecule (common in prokaryotic cells)
or a number of DNA molecules (common
in eukaryotic cells)
DNA molecules in a cell are packaged
into chromosomes
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FIGURE 12.3
20 m
Eukaryotic chromosomes consist of chromatin,
a complex of DNA and protein that condenses
during cell division
Every eukaryotic species has a characteristic
number of chromosomes in each cell nucleus
Somatic cells (nonreproductive cells) have two
sets of chromosomes
Gametes (reproductive cells: sperm and eggs)
have half as many chromosomes as somatic
cells
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DISTRIBUTION OF CHROMOSOMES
DURING EUKARYOTIC CELL DIVISION
Before cell division, DNA is replicated and the
chromosomes condense
Each duplicated chromosome has two sister
chromatids (joined copies of the original
chromosome), which separate during cell
division
The centromere is the narrow “waist” of the
duplicated chromosome, where the two
chromatids are most closely attached
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FIGURE 12.4
Sister
chromatids
Centromere
0.5 m
During cell division, the two sister
chromatids of each duplicated
chromosome separate and move into
two nuclei
Once separate, the chromatids are
called chromosomes
© 2011 Pearson Education, Inc.
FIGURE 12.5-1
Chromosomes
1
Chromosomal
DNA molecules
Centromere
Chromosome
arm
FIGURE 12.5-2
Chromosomes
1
Chromosomal
DNA molecules
Centromere
Chromosome
arm
Chromosome duplication
(including DNA replication)
and condensation
2
Sister
chromatids
FIGURE 12.5-3
Chromosomes
1
Chromosomal
DNA molecules
Centromere
Chromosome
arm
Chromosome duplication
(including DNA replication)
and condensation
2
Sister
chromatids
Separation of sister
chromatids into
two chromosomes
3
Eukaryotic cell division consists of
 Mitosis, the division of the genetic material
in the nucleus
 Cytokinesis, the division of the cytoplasm
Gametes are produced by a variation of cell
division called meiosis
Meiosis yields nonidentical daughter cells that
have only one set of chromosomes, half as
many as the parent cell
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PHASES OF THE CELL CYCLE
The cell cycle consists of
 Mitotic (M) phase (mitosis and cytokinesis)
 Interphase (cell growth and copying of
chromosomes in preparation for cell
division)
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Interphase (about 90% of the cell cycle)
can be divided into subphases
 G 1 phase (“first gap”)
 S phase (“synthesis”)
 G 2 phase (“second gap”)
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FIGURE 12.6
INTERPHASE
G1
S
(DNA synthesis)
G2
Mitosis is conventionally divided into five
phases
 Prophase
 Prometaphase
 Metaphase
 Anaphase
 Telophase
Cytokinesis overlaps the latter stages of
mitosis
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BioFlix: Mitosis
© 2011 Pearson Education, Inc.
10 m
FIGURE 12.7
G2 of Interphase
Centrosomes
(with centriole pairs)
Nucleolus
Chromatin
(duplicated)
Nuclear
envelope
Plasma
membrane
Prophase
Early mitotic
spindle
Aster
Centromere
Chromosome, consisting
of two sister chromatids
Prometaphase
Fragments
of nuclear
envelope
Kinetochore
Metaphase
Nonkinetochore
microtubules
Kinetochore
microtubule
Anaphase
Metaphase
plate
Spindle
Centrosome at
one spindle pole
Telophase and Cytokinesis
Cleavage
furrow
Daughter
chromosomes
Nuclear
envelope
forming
Nucleolus
forming
FIGURE 12.7A
G2 of Interphase
Centrosomes
(with centriole
pairs)
Chromatin
(duplicated)
Prophase
Early mitotic
spindle
Plasma
membrane
Nucleolus
Nuclear
envelope
Aster
Centromere
Chromosome, consisting
of two sister chromatids
Prometaphase
Fragments
of nuclear
envelope
Kinetochore
Nonkinetochore
microtubules
Kinetochore
microtubule
FIGURE 12.7B
Metaphase
Anaphase
Metaphase
plate
Spindle
Centrosome at
one spindle pole
Telophase and Cytokinesis
Cleavage
furrow
Daughter
chromosomes
Nuclear
envelope
forming
Nucleolus
forming
10 m
FIGURE 12.7C
G2 of Interphase
Prophase
Prometaphase
10 m
FIGURE 12.7D
Metaphase
Anaphase
Telophase and Cytokinesis
THE MITOTIC SPINDLE: A CLOSER
LOOK
The mitotic spindle is a structure made of
microtubules that controls chromosome
movement during mitosis
In animal cells, assembly of spindle
microtubules begins in the centrosome,
the microtubule organizing center (MTOC)
The centrosome replicates during
interphase, forming two centrosomes that
migrate to opposite ends of the cell
during prophase and prometaphase
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An aster (a radial array of short
microtubules) extends from each
centrosome
The spindle includes the centrosomes,
the spindle microtubules, and the asters
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 During prometaphase, some spindle
microtubules attach to the kinetochores of
chromosomes and begin to move the
chromosomes
 Kinetochores are protein complexes associated
with centromeres
 At metaphase, the chromosomes are all lined up
at the metaphase plate, an imaginary structure
at the midway point between the spindle’s two
poles
© 2011 Pearson Education, Inc.
FIGURE 12.8
Aster
Centrosome
Sister
chromatids
Metaphase
plate
(imaginary)
Microtubules
Chromosomes
Kinetochores
Centrosome
1 m
Overlapping
nonkinetochore
microtubules
Kinetochore
microtubules
0.5 m
In anaphase, sister chromatids separate
and move along the kinetochore
microtubules toward opposite ends of
the cell
The microtubules shorten by
depolymerizing at their kinetochore
ends
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FIGURE 12.9
EXPERIMENT
Kinetochore
Spindle
pole
Mark
RESULTS
CONCLUSION
Microtubule
Chromosome
movement
Motor protein
Chromosome
Kinetochore
Tubulin
subunits
In telophase, genetically identical
daughter nuclei form at opposite
ends of the cell
Cytokinesis begins during anaphase
or telophase and the spindle
eventually disassembles
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CY TOKINESIS: A CLOSER LOOK
In animal cells, cytokinesis occurs by a
process known as cleavage, forming a
cleavage furrow
In plant cells, a cell plate forms during
cytokinesis
© 2011 Pearson Education, Inc.
Animation: Cytokinesis
Right-click slide / select ”Play”
© 2011 Pearson Education, Inc.
Animation: Animal Mitosis
Right-click slide / select ”Play”
© 2011 Pearson Education, Inc.
Animation: Sea Urchin (Time Lapse)
Right-click slide / select ”Play”
© 2011 Pearson Education, Inc.
FIGURE 12.10
(a) Cleavage of an animal cell (SEM)
100 m
Cleavage furrow
Contractile ring of
microfilaments
(b) Cell plate formation in a plant cell (TEM)
Vesicles
forming
cell plate
Wall of parent cell
Cell plate
1 m
New cell wall
Daughter cells
Daughter cells
F I G U R E 1 2 . 11
Nucleus
Chromatin
condensing
Nucleolus
1 Prophase
Chromosomes
2 Prometaphase 3 Metaphase
Cell plate
4 Anaphase
10 m
5 Telophase
F I G U R E 1 2 . 11 A
Nucleus
Chromatin
condensing
Nucleolus
10 m
1 Prophase
BINARY FISSION IN BACTERIA
Prokaryotes (bacteria and archaea)
reproduce by a type of cell division called
binary fission
In binary fission, the chromosome
replicates (beginning at the origin of
replication), and the two daughter
chromosomes actively move apart
The plasma membrane pinches inward,
dividing the cell into two
© 2011 Pearson Education, Inc.
FIGURE 12.12-4
Origin of
replication
E. coli cell
1 Chromosome
replication
begins.
2 Replication
continues.
3 Replication
finishes.
4 Two daughter
cells result.
Cell wall
Plasma membrane
Bacterial chromosome
Two copies
of origin
Origin
Origin
THE EVOLUTION OF MITOSIS
Since prokaryotes evolved before eukaryotes,
mitosis probably evolved from binary fission
Certain protists exhibit types of cell division
that seem intermediate between binary fission
and mitosis
© 2011 Pearson Education, Inc.
FIGURE 12.13
(a) Bacteria
Bacterial
chromosome
Chromosomes
(b) Dinoflagellates
Microtubules
Intact nuclear
envelope
Kinetochore
microtubule
(c) Diatoms and
some yeasts
Intact nuclear
envelope
Kinetochore
microtubule
(d) Most eukaryotes
Fragments of
nuclear envelope
FIGURE 12.14
EXPERIMENT
Experiment 1
S
G1
S
S
Experiment 2
M
G1
RESULTS
When a cell in the S
phase was fused
with a cell in G1,
the G1 nucleus
immediately entered
the S phase—DNA
was synthesized.
M
M
When a cell in the
M phase was fused with
a cell in G1, the G1
nucleus immediately
began mitosis—a spindle
formed and chromatin
condensed, even though
the chromosome had not
been duplicated.
THE CELL CYCLE CONTROL SYSTEM
The events of the cell cycle are directed
by a cell cycle control system
The cell cycle control system is
regulated by both internal and external
controls
The clock has specific checkpoints
where the cell cycle stops until a goahead signal is received
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THE MAP SIGNAL TRANSDUCTION
PATHWAY AND P53 PROTEIN
 When a mitogen binds to a RTK, the Ras G-protein activates
MAK cascade (from pg.81).
 This activates a transcription factor that signals the
expression of p53 gene to p53 protein
 P53 protein checks for damaged DNA and directs enzymes to
repair the DNA
 If the DNA cannot be repaired, the cell is signaled to die –
apoptosis
 If p53 gene is mutated or if there is damage to any protein on
the MAP pathway, the p53 protein will be altered or non exsistent.
 This allows cells to divide without proper DNA structure.
FIGURE 12.15
G1 checkpoint
Control
system
G1
M
G2
M checkpoint
G2 checkpoint
S
the G 1 checkpoint is the most important
If a cell receives a go-ahead signal at
the G 1 checkpoint, it will usually
complete the S, G 2, and M phases and
divide
If the cell does not receive the go-ahead
signal, it will exit the cycle, switching
into a nondividing state called the G 0
phase
© 2011 Pearson Education, Inc.
FIGURE 12.16
G0
G1 checkpoint
G1
(a) Cell receives a go-ahead
signal.
G1
(b) Cell does not receive a
go-ahead signal.
THE CELL CYCLE CLOCK: CYCLINS AND
CYCLIN-DEPENDENT KINASES
 Two types of regulatory proteins are involved in cell cycle
control: cyclins and cyclin-dependent kinases (Cdks)
 Cdks activity fluctuates during the cell cycle because it is
controled by cyclins, so named because their concentrations
vary with the cell cycle
 MPF (maturation-promoting factor) is a cyclin-Cdk complex that
triggers a cell’s passage past the G 2 checkpoint into the M
phase
© 2011 Pearson Education, Inc.
FIGURE 12.17
M
G1
S G2
M
G1 S
G2
M
G1
MPF activity
Cyclin
concentration
Time
(a) Fluctuation of MPF activity and cyclin concentration
during the cell cycle
Cdk
Degraded
cyclin
Cyclin is
degraded
G2
Cdk
checkpoint
MPF
Cyclin
(b) Molecular mechanisms that help regulate the cell cycle
STOP AND GO SIGNS: INTERNAL AND
EXTERNAL SIGNALS AT THE CHECKPOINTS
 An example of an internal signal is that kinetochores not attached
to spindle microtubules send a molecular signal that delays
anaphase
 Platelet-derived growth factor (PDGF) stimulates the division of
human fibroblast cells in culture
 External growth factors can activate a kinase cascade,
which activates the RAS protein
 Ras activate a MAK cascade which stimulates
transcription factors
 These land on p53 gene and synthesize the p53 protein
 P53 protein checks DNA in the cell cycle
 If any part of the cascade is mutated, the cell cycle goes
unchecked.
 Video and cyclin/CDK video
© 2011 Pearson Education, Inc.
FIGURE 12.18
1 A sample of human
connective tissue is
cut up into small
pieces.
Scalpels
Petri
dish
2 Enzymes digest
the extracellular
matrix, resulting in
a suspension of
free fibroblasts.
3 Cells are transferred to
culture vessels.
Without PDGF
4 PDGF is added
to half the
vessels.
With PDGF
10 m
FIGURE 12.18A
10 m
EXTERNAL FEATURES THAT AFFECT CELL
DIVISON
 density -dependent inhibition, in which crowded cells stop
dividing
 Most animal cells also exhibit anchorage dependence, in
which they must be attached to a substratum in order to
divide
 Cancer cells exhibit neither density -dependent inhibition nor
anchorage dependence
© 2011 Pearson Education, Inc.
FIGURE 12.19
Anchorage dependence
Density-dependent inhibition
Density-dependent inhibition
20 m
20 m
(a) Normal mammalian cells
(b) Cancer cells
LOSS OF CELL CYCLE CONTROLS IN CANCER
CELLS
 Cancer cells do not respond normally to the body’s control
mechanisms
 Cancer cells may not need growth factors to grow and divide
 They may make their own growth factor
 They may convey a growth factor’s signal without the presence of
the growth factor
 They may have an abnormal cell cycle control system
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 A normal cell is converted to a cancerous cell by a process
called transformation
 Cancer cells that are not eliminated by the immune system
form tumors, masses of abnormal cells within otherwise
normal tissue
 If abnormal cells remain only at the original site, the lump is
called a benign tumor
 Malignant tumors invade surrounding tissues and can
metastasize, exporting cancer cells to other parts of the body,
where they may form additional tumors
© 2011 Pearson Education, Inc.
FIGURE 12.20
Tumor
Lymph
vessel
Blood
vessel
Glandular
tissue
Cancer
cell
1 A tumor grows
from a single
cancer cell.
Metastatic
tumor
2 Cancer
cells invade
neighboring
tissue.
3 Cancer cells spread
through lymph and
blood vessels to
other parts of the
body.
4 Cancer cells
may survive
and establish
a new tumor
in another part
of the body.
FIGURE 12.UN01
P
G1
S
Cytokinesis
Mitosis
G2
MITOTIC (M) PHASE
Prophase
Telophase and
Cytokinesis
Prometaphase
Anaphase
Metaphase
FIGURE 12.UN02
FIGURE 12.UN03
FIGURE 12.UN04
FIGURE 12.UN05
FIGURE 12.UN06