1. Nonkinetochore microtubules from opposite poles overlap and push against each other, elongating the cell
In telophase, genetically identical daughter nuclei form at opposite ends of the cell
For the Cell Biology Video Microtubules in Cell Division, go to Animation and Video Files.
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2. Cytokinesis: 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
Animation: Cytokinesis
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3. Video: Sea Urchin (Time Lapse)
Video: Animal Mitosis
Video: Sea Urchin (Time Lapse)
For the Cell Biology Video Nuclear Envelope Formation, go to Animation and Video Files.
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

4. Figure 12.9 Cytokinesis in animal and plant cells
Vesicles
forming
cell plate
Wall of
parent cell
1 µm
100 µm
Cleavage furrow
Cell plate
New cell wall
Figure 12.9 Cytokinesis in animal and plant cells
Contractile ring of
microfilaments
Daughter cells
Daughter cells
(a) Cleavage of an animal cell (SEM)
(b) Cell plate formation in a plant cell (TEM)

5. 10 µm Fig. 12-10 Nucleus Chromatin condensing Nucleolus Chromosomes
Cell plate
Figure Mitosis in a plant cell 1
Prophase 2
Prometaphase 3
Metaphase 4
Anaphase 5
Telophase

6. Binary Fission
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
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7. Cell wall Origin of replication Plasma membrane E. coli cell Bacterial
Fig
Cell wall
Origin of
replication
Plasma
membrane
E. coli cell
Bacterial
chromosome
Two copies
of origin
Origin
Origin
Figure Bacterial cell division by binary fission

8. 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
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9. Fig
Bacterial
chromosome
(a) Bacteria
Chromosomes
Microtubules
Intact nuclear
envelope
(b) Dinoflagellates
Kinetochore
microtubule
Intact nuclear
envelope
Figure A hypothetical sequence for the evolution of mitosis
(c) Diatoms and yeasts
Kinetochore
microtubule
Fragments of
nuclear envelope
(d) Most eukaryotes

10. The frequency of cell division varies with the type of cell
Concept 12.3: The eukaryotic cell cycle is regulated by a molecular control system
The frequency of cell division varies with the type of cell
These cell cycle differences result from regulation at the molecular level
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11. Evidence for Cytoplasmic Signals
The cell cycle appears to be driven by specific chemical signals present in the cytoplasm
Some evidence for this hypothesis comes from experiments in which cultured mammalian cells at different phases of the cell cycle were fused to form a single cell with two nuclei
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12. EXPERIMENT RESULTS Experiment 1 Experiment 2 S G1 M G1 S S M M
Fig
EXPERIMENT
Experiment 1
Experiment 2 S G1 M G1
RESULTS S S M M
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.
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.
Figure Do molecular signals in the cytoplasm regulate the cell cycle?

13. The Cell Cycle Control System
The sequential events of the cell cycle are directed by a distinct cell cycle control system, which is similar to a clock
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 go-ahead signal is received
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14. G1 checkpoint Control system S G1 G2 M M checkpoint G2 checkpoint
Fig
G1 checkpoint
Control
system S G1 G2 M
Figure Mechanical analogy for the cell cycle control system
M checkpoint
G2 checkpoint

15. For many cells, the G1 checkpoint seems to be the most important one
If a cell receives a go-ahead signal at the G1 checkpoint, it will usually complete the S, G2, 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 G0 phase
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16. G0 G1 G1 G1 checkpoint (b) Cell does not receive a go-ahead signal
Fig G0
G1 checkpoint G1 G1
Figure The G1 checkpoint
Cell receives a go-ahead
signal
(b) Cell does not receive a
go-ahead signal

17. 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)
The activity of cyclins and Cdks fluctuates during the cell cycle
MPF (maturation-promoting factor) is a cyclin-Cdk complex that triggers a cell’s passage past the G2 checkpoint into the M phase
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18. Fig
RESULTS 5 30 4 20 3
% of dividing cells (– )
Protein kinase activity (– ) 2 10 1
Figure How does the activity of a protein kinase essential for mitosis vary during the cell cycle?
100
200
300
400
500
Time (min)

19. M G1 S G2 M G1 S G2 M G1 Fig. 12-17 MPF activity Cyclin concentration
Time
(a) Fluctuation of MPF activity and cyclin concentration during
the cell cycle G1 S
Cdk
Figure Molecular control of the cell cycle at the G2 checkpoint
Cyclin accumulation M
Degraded
cyclin G2 G2
Cdk
Cyclin is
degraded
checkpoint
Cyclin
MPF
(b) Molecular mechanisms that help regulate the cell cycle

20. 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
Some external signals are growth factors, proteins released by certain cells that stimulate other cells to divide
For example, platelet-derived growth factor (PDGF) stimulates the division of human fibroblast cells in culture
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21. Scalpels Petri plate Without PDGF cells fail to divide With PDGF
Fig
Scalpels
Petri
plate
Without PDGF
cells fail to divide
Figure The effect of a growth factor on cell division
With PDGF
cells prolifer-
ate
Cultured fibroblasts
10 µm

22. Another example of external signals is 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
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23. Density-dependent inhibition
Fig
Anchorage dependence
Density-dependent inhibition
Density-dependent inhibition
Figure Density-dependent inhibition and anchorage dependence of cell division
25 µm
25 µm
(a) Normal mammalian cells
(b) Cancer cells

24. Cancer cells exhibit neither density-dependent inhibition nor anchorage dependence
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25. 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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26. A normal cell is converted to a cancerous cell by a process called transformation
Cancer cells form tumors, masses of abnormal cells within otherwise normal tissue
If abnormal cells remain 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 secondary tumors
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

27. Lymph vessel Tumor Blood vessel Cancer cell Glandular tissue
Fig
Lymph
vessel
Tumor
Blood
vessel
Cancer
cell
Glandular
tissue
Metastatic
tumor 1
A tumor grows
from a single
cancer cell. 2
Cancer cells
invade neigh-
boring tissue. 3
Cancer cells spread
to other parts of
the body. 4
Cancer cells may
survive and
establish a new
tumor in another
part of the body.
Figure The growth and metastasis of a malignant breast tumor

28. The Immortal Cells of Henrietta Lacks
This new species is Helacyton gartleri; it is the cell line cultured from the diseased cervix of Henrietta Lacks.
The culture was created by George and Margaret Gey in They were researchers who were sent a sample of Henrietta Lack’s’ tumor. They had wanted to understand cancer, but were frustrated by, among other things, by the inability to maintain a culture of human cells in the laboratory. Inevitably, cells taken for culture would weaken and die. They had been trying for 20 years grow human cells in vitro without success. Until Henrietta Lacks. Her cells were found to be amazingly viable, and the Hela cell line was underway. A little too underway, as it turned out as she died a mere 9 months later from cancer that had spread throughout her body
Modern DNA techniques can distinguish Hela cells from other cell lines, and they can also show that Henrietta Lacks cancer was initially cervical cancer, probably the result of human papilloma virus infection.
The cells were cultured without her knowledge or consent, and her family remained ignorant of her role until Henrietta Lacks's contribution to science is now more openly acknowledged, but still little known.