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Chapter 12

The Cell Cycle

PowerPoint Lectures for
Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Overview: The Key Roles of Cell Division
• The continuity of life
– Is based upon the reproduction of cells, or cell
division

Figure 12.1
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Unicellular organisms
– Reproduce by cell division
100 µm

(a) Reproduction. An amoeba,
a single-celled eukaryote, is
dividing into two cells. Each
new cell will be an individual
Figure 12.2 A
organism (LM).

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Multicellular organisms depend on cell division
for
– Development from a fertilized cell
– Growth
– Repair

200 µm

20 µm

(b) Growth and development.
(c) Tissue renewal. These dividing
This micrograph shows a
bone marrow cells (arrow) will
sand dollar embryo shortly
give rise to new blood cells (LM).
after the fertilized egg divided,
Figure 12.2 B, C forming two cells (LM).
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• The cell division process
– Is an integral part of the cell cycle

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Concept 12.1: Cell division results in
genetically identical daughter cells
• Cells duplicate their genetic material
– Before they divide, ensuring that each
daughter cell receives an exact copy of the
genetic material, DNA

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Cellular Organization of the Genetic Material
• A cell’s endowment of DNA, its genetic
information
– Is called its genome

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• The DNA molecules in a cell
– Are packaged into chromosomes

Figure 12.3
50 µm
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Eukaryotic chromosomes
– Consist of chromatin, a complex of DNA and
protein that condenses during cell division

• In animals
– Somatic cells have two sets of chromosomes
– Gametes have one set of chromosomes

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Distribution of Chromosomes During Cell Division
• In preparation for cell division
– DNA is replicated and the chromosomes
condense

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Each duplicated chromosome
– Has two sister chromatids, which separate
during cell division
0.5 µm
A eukaryotic cell has multiple
chromosomes, one of which is
represented here. Before
duplication, each chromosome
has a single DNA molecule.

Once duplicated, a chromosome
consists of two sister chromatids
connected at the centromere. Each
chromatid contains a copy of the
DNA molecule.

Mechanical processes separate
the sister chromatids into two
chromosomes and distribute
them to two daughter cells.

Figure 12.4
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Chromosome
duplication
(including DNA
synthesis)
Centromere

Separation
of sister
chromatids

Centromeres

Sister
chromatids

Sister chromatids
• Eukaryotic cell division consists of
– Mitosis, the division of the nucleus
– Cytokinesis, the division of the cytoplasm

• In meiosis
– Sex cells are produced after a reduction in
chromosome number

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Concept 12.2: The mitotic phase alternates
with interphase in the cell cycle
• A labeled probe can reveal patterns of gene
expression in different kinds of cells

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Phases of the Cell Cycle
• The cell cycle consists of
– The mitotic phase
– Interphase
INTERPHASE

S
(DNA synthesis)

C
M yto
ito ki
si ne
s si

s

G1

MI
(M TOT
) P IC
HA
SE

Figure 12.5
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

G2
• Interphase can be divided into subphases
– G1 phase
– S phase
– G2 phase

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• The mitotic phase
– Is made up of mitosis and cytokinesis

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Mitosis consists of five distinct phases
– Prophase
– Prometaphase

G2 OF
INTERPHASE
Centrosomes
Chromatin
(with centriole pairs)
(duplicated)

Figure 12.6

Nucleolus

Nuclear
Plasma
envelope membrane

PROPHASE
Early mitotic
spindle

Aster
Centromere

Chromosome, consisting
of two sister chromatids

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

PROMETAPHASE
Fragments Kinetochore
of nuclear
envelope
Nonkinetochore
microtubules

Kinetochore
microtubule
– Metaphase
– Anaphase
– Telophase

METAPHASE

ANAPHASE

Metaphase
plate

Figure 12.6

Spindle

Centrosome at Daughter
one spindle pole chromosomes

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

TELOPHASE AND CYTOKINESIS
Cleavage
furrow

Nuclear
envelope
forming

Nucleolus
forming
The Mitotic Spindle: A Closer Look
• The mitotic spindle
– Is an apparatus of microtubules that controls
chromosome movement during mitosis

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• The spindle arises from the centrosomes
– And includes spindle microtubules and asters

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Some spindle microtubules
–

Attach to the kinetochores of chromosomes and move the
chromosomes to the metaphase plate
Aster
Sister
chromatids

Centrosome
Metaphase
Plate
Kinetochores

Overlapping
nonkinetochore
microtubules
Kinetochores
microtubules
Microtubules

0.5 µm

Figure 12.7 Centrosome
1 µm
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Chromosomes
• In anaphase, sister chromatids separate
– And move along the kinetochore microtubules
toward opposite ends of the cell
EXPERIMENT

1 The microtubules of a cell in early anaphase were labeled with a fluorescent dye
that glows in the microscope (yellow).

Kinetochore

Spindle
pole

Figure 12.8
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Nonkinetechore 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

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Cytokinesis: A Closer Look
• In animal cells
– Cytokinesis occurs by a process known as
cleavage, forming a cleavage furrow

Cleavage furrow

Contractile ring of
microfilaments

Figure 12.9 A

100 µm

Daughter cells

(a) Cleavage of an animal cell (SEM)

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• In plant cells, during cytokinesis
– A cell plate forms

Vesicles
forming
cell plate

1 µm
Wall of
patent cell Cell plate New cell wall

Daughter cells
Figure 12.9 B (b) Cell plate formation in a plant cell (SEM)
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Mitosis in a plant cell
Chromatine
Nucleus
Nucleolus condensing

Chromosome

Metaphase. The
Anaphase. The
2 Prometaphase.
3
5
1 Prophase.
spindle is complete, 4 chromatids of each
The chromatin
We now see discrete
and the chromosomes, chromosome have
is condensing.
chromosomes; each
attached to microtubules separated, and the
The nucleolus is
consists of two
at their kinetochores,
beginning to
identical sister
daughter chromosomes
are all at the metaphase are moving to the ends
disappear.
chromatids. Later
plate.
Although not
in prometaphase, the
of cell as their
yet visible
nuclear envelop will
kinetochore
in the micrograph,
fragment.
microtubles shorten.
the mitotic spindle is
staring to from.

Figure 12.10
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Telophase. Daughter
nuclei are forming.
Meanwhile, cytokinesis
has started: The cell
plate, which will
divided the cytoplasm
in two, is growing
toward the perimeter
of the parent cell.
Binary Fission
• Prokaryotes (bacteria)
– Reproduce by a type of cell division called
binary fission

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• In binary fission
– The bacterial chromosome replicates
– The two daughter chromosomes actively move apart
Origin of
replication

Cell wall

E. coli cell
Chromosome replication
1 begins.
Soon thereafter, one copy of
the origin moves rapidly toward
the other end of the cell.
2 Replication continues. One copy of
the origin is now at each end of
the cell.

3 Replication finishes. The plasma
membrane grows inward, and
new cell wall is deposited.

Figure 12.11 4 Two daughter cells result.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Two copies
of origin

Origin

Plasma
Membrane
Bacterial
Chromosom
e

Origin
The Evolution of Mitosis
• Since prokaryotes preceded eukaryotes by
billions of years
– It is likely that mitosis evolved from bacterial
cell division

• Certain protists
– Exhibit types of cell division that seem
intermediate between binary fission and
mitosis carried out by most eukaryotic cells

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• A hypothetical sequence for the evolution of mitosis
(a) Prokaryotes. During binary fission, the origins of the
daughter chromosomes move to opposite ends of the
cell. The mechanism is not fully understood, but
proteins may anchor the daughter chromosomes to
specific sites on the plasma membrane.
(b) Dinoflagellates. In unicellular protists called
dinoflagellates, the nuclear envelope remains intact
during cell division, and the chromosomes attach to the
nuclear envelope. Microtubules pass through the
nucleus inside cytoplasmic tunnels, reinforcing the
spatial orientation of the nucleus, which then divides in a
fission process reminiscent of bacterial division.
(c) Diatoms. In another group of unicellular protists, the
diatoms, the nuclear envelope also remains intact
during cell division. But in these organisms, the
microtubules form a spindle within the nucleus.
Microtubules separate the chromosomes, and the
nucleus splits into two daughter nuclei.
(d) Most eukaryotes. In most other eukaryotes,
including plants and animals, the spindle forms
outside the nucleus, and the nuclear envelope
breaks down during mitosis. Microtubules
separate the chromosomes, and the nuclear
envelope then re-forms.

Figure 12.12 A-D
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Bacterial
chromosome
Chromosomes
Microtubules
Intact nuclear
envelope
Kinetochore
microtubules
Intact nuclear
envelope
Kinetochore
microtubules
Centrosome
Fragments of
nuclear envelope
• Concept 12.3: The 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

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Evidence for Cytoplasmic Signals
• Molecules present in the cytoplasm
– Regulate progress through the cell cycle
EXPERIMENTS In each experiment, cultured mammalian cells at two different phases of the cell cycle were induced to fuse.

Experiment 2

Experiment 1

S

G1

M

S

M

G1

RESULTS

S

When a cell in the S
phase was fused with
a cell in G1, the G1 cell
immediately entered the
S phase—DNA was
synthesized.

M

When a cell in the M phase
was fused with a cell in G1, the
G1 cell immediately began mitosis—
a spindle formed and chromatin
condensed, even though the
chromosome had not been duplicated.

CONCLUSION The results of fusing cells at two different phases of the cell cycle suggest that molecules present in the

Figure 12.13 A, B

cytoplasm of cells in the S or M phase control the progression of phases.

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
G1 checkpoint

Control
system
G1

M

G2

M checkpoint
Figure 12.14
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

G2 checkpoint

S
• The clock has specific checkpoints
– Where the cell cycle stops until a go-ahead signal is
received

G0
G1 checkpoint

G1
(a) If a cell receives a go-ahead signal at
the G1 checkpoint, the cell continues

Figure 12.15 A, B

on in the cell cycle.

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

G1
(b) If a cell does not receive a go-ahead
signal at the G1checkpoint, the cell
exits the cell cycle and goes into G0, a
nondividing state.
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)

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• The activity of cyclins and Cdks
– Fluctuates during the cell cycle
(a) Fluctuation of MPF activity and
cyclin concentration during
the cell cycle

M

G1 S G2 M

G1 S G2 M

MPF activity
Cyclin

Time
(b) Molecular mechanisms that
help regulate the cell cycle

1 Synthesis of cyclin begins in late S
phase and continues through G2.
Because cyclin is protected from
degradation during this stage, it
accumulates.

G

4 During anaphase, the cyclin component
of MPF is degraded, terminating the M
phase. The cell enters the G1 phase.

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

2

G

Degraded
Cyclin
Cyclin is
degraded

Figure 12.16 A, B

S

Cdk
M

the cell favor degradation
of cyclin, and the Cdk
component of MPF is
recycled.

1

5 During G1, conditions in

G2
Cdk
checkpoint

MPF

Cyclin

2 Accumulated cyclin molecules

combine with recycled Cdk molecules, producing enough molecules
of MPF to pass the G2 checkpoint and
initiate the events of mitosis.

3 MPF promotes mitosis by phosphorylating
various proteins. MPF‘s activity peaks during
metaphase.
Stop and Go Signs: Internal and External Signals at
the Checkpoints
• Both internal and external signals
– Control the cell cycle checkpoints

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Growth factors
– Stimulate other cells to divide
EXPERIMENT

Scalpels

1 A sample of connective tissue was cut up

into small pieces.

Petri
plate
2 Enzymes were used to digest the extracellular matrix,

resulting in a suspension of free fibroblast cells.

3 Cells were transferred to sterile culture vessels

Figure 12.17

containing a basic growth medium consisting of
glucose, amino acids, salts, and antibiotics (as a
Without PDGF
precaution against bacterial growth). PDGF was
added to half the vessels. The culture vessels
were incubated at 37°C.
With PDGF

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• In density-dependent inhibition
– Crowded cells stop dividing
• Most animal cells exhibit anchorage dependence
– In which they must be attached to a substratum to
divide
(a) Normal mammalian cells. The
availability of nutrients, growth
factors, and a substratum for
attachment limits cell
density to a single layer.

Cells anchor to dish surface and
divide (anchorage dependence).
When cells have formed a complete single layer,
they stop dividing
(density-dependent inhibition).

If some cells are scraped away, the remaining cells
divide to fill the gap and then stop (densitydependent inhibition).

Figure 12.18 A
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

25 µm
• Cancer cells
– Exhibit neither density-dependent inhibition nor
anchorage dependence
Cancer cells do not exhibit
anchorage dependence or
density-dependent inhibition.
(b) Cancer cells. Cancer cells usually
continue to divide well beyond a
single layer, forming a clump of
overlapping cells.

Figure 12.18 B
25 µm

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Loss of Cell Cycle Controls in Cancer Cells
• Cancer cells
– Do not respond normally to the body’s control
mechanisms
– Form tumors

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Malignant tumors invade surrounding tissues
and can metastasize
– Exporting cancer cells to other parts of the
body where they may form secondary tumors

Lymph
vessel

Tumor

Blood
vessel

Glandular
tissue
1 A tumor grows from a
single cancer cell.

Cancer cell
2 Cancer cells invade
neighboring tissue.

Figure 12.19
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

3 Cancer cells spread
through lymph and
blood vessels to
other parts of the body.

Metastatic
Tumor
4 A small percentage of
cancer cells may survive
and establish a new tumor
in another part of the body.

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12 cellcycle text

  • 1. Chapter 12 The Cell Cycle PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 2. • Overview: The Key Roles of Cell Division • The continuity of life – Is based upon the reproduction of cells, or cell division Figure 12.1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 3. • Unicellular organisms – Reproduce by cell division 100 µm (a) Reproduction. An amoeba, a single-celled eukaryote, is dividing into two cells. Each new cell will be an individual Figure 12.2 A organism (LM). Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 4. • Multicellular organisms depend on cell division for – Development from a fertilized cell – Growth – Repair 200 µm 20 µm (b) Growth and development. (c) Tissue renewal. These dividing This micrograph shows a bone marrow cells (arrow) will sand dollar embryo shortly give rise to new blood cells (LM). after the fertilized egg divided, Figure 12.2 B, C forming two cells (LM). Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 5. • The cell division process – Is an integral part of the cell cycle Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 6. • Concept 12.1: Cell division results in genetically identical daughter cells • Cells duplicate their genetic material – Before they divide, ensuring that each daughter cell receives an exact copy of the genetic material, DNA Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 7. Cellular Organization of the Genetic Material • A cell’s endowment of DNA, its genetic information – Is called its genome Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 8. • The DNA molecules in a cell – Are packaged into chromosomes Figure 12.3 50 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 9. • Eukaryotic chromosomes – Consist of chromatin, a complex of DNA and protein that condenses during cell division • In animals – Somatic cells have two sets of chromosomes – Gametes have one set of chromosomes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 10. Distribution of Chromosomes During Cell Division • In preparation for cell division – DNA is replicated and the chromosomes condense Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 11. • Each duplicated chromosome – Has two sister chromatids, which separate during cell division 0.5 µm A eukaryotic cell has multiple chromosomes, one of which is represented here. Before duplication, each chromosome has a single DNA molecule. Once duplicated, a chromosome consists of two sister chromatids connected at the centromere. Each chromatid contains a copy of the DNA molecule. Mechanical processes separate the sister chromatids into two chromosomes and distribute them to two daughter cells. Figure 12.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Chromosome duplication (including DNA synthesis) Centromere Separation of sister chromatids Centromeres Sister chromatids Sister chromatids
  • 12. • Eukaryotic cell division consists of – Mitosis, the division of the nucleus – Cytokinesis, the division of the cytoplasm • In meiosis – Sex cells are produced after a reduction in chromosome number Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 13. • Concept 12.2: The mitotic phase alternates with interphase in the cell cycle • A labeled probe can reveal patterns of gene expression in different kinds of cells Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 14. Phases of the Cell Cycle • The cell cycle consists of – The mitotic phase – Interphase INTERPHASE S (DNA synthesis) C M yto ito ki si ne s si s G1 MI (M TOT ) P IC HA SE Figure 12.5 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings G2
  • 15. • Interphase can be divided into subphases – G1 phase – S phase – G2 phase Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 16. • The mitotic phase – Is made up of mitosis and cytokinesis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 17. • Mitosis consists of five distinct phases – Prophase – Prometaphase G2 OF INTERPHASE Centrosomes Chromatin (with centriole pairs) (duplicated) Figure 12.6 Nucleolus Nuclear Plasma envelope membrane PROPHASE Early mitotic spindle Aster Centromere Chromosome, consisting of two sister chromatids Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings PROMETAPHASE Fragments Kinetochore of nuclear envelope Nonkinetochore microtubules Kinetochore microtubule
  • 18. – Metaphase – Anaphase – Telophase METAPHASE ANAPHASE Metaphase plate Figure 12.6 Spindle Centrosome at Daughter one spindle pole chromosomes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings TELOPHASE AND CYTOKINESIS Cleavage furrow Nuclear envelope forming Nucleolus forming
  • 19. The Mitotic Spindle: A Closer Look • The mitotic spindle – Is an apparatus of microtubules that controls chromosome movement during mitosis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 20. • The spindle arises from the centrosomes – And includes spindle microtubules and asters Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 21. • Some spindle microtubules – Attach to the kinetochores of chromosomes and move the chromosomes to the metaphase plate Aster Sister chromatids Centrosome Metaphase Plate Kinetochores Overlapping nonkinetochore microtubules Kinetochores microtubules Microtubules 0.5 µm Figure 12.7 Centrosome 1 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Chromosomes
  • 22. • In anaphase, sister chromatids separate – And move along the kinetochore microtubules toward opposite ends of the cell EXPERIMENT 1 The microtubules of a cell in early anaphase were labeled with a fluorescent dye that glows in the microscope (yellow). Kinetochore Spindle pole Figure 12.8 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 23. • Nonkinetechore 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 24. Cytokinesis: A Closer Look • In animal cells – Cytokinesis occurs by a process known as cleavage, forming a cleavage furrow Cleavage furrow Contractile ring of microfilaments Figure 12.9 A 100 µm Daughter cells (a) Cleavage of an animal cell (SEM) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 25. • In plant cells, during cytokinesis – A cell plate forms Vesicles forming cell plate 1 µm Wall of patent cell Cell plate New cell wall Daughter cells Figure 12.9 B (b) Cell plate formation in a plant cell (SEM) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 26. • Mitosis in a plant cell Chromatine Nucleus Nucleolus condensing Chromosome Metaphase. The Anaphase. The 2 Prometaphase. 3 5 1 Prophase. spindle is complete, 4 chromatids of each The chromatin We now see discrete and the chromosomes, chromosome have is condensing. chromosomes; each attached to microtubules separated, and the The nucleolus is consists of two at their kinetochores, beginning to identical sister daughter chromosomes are all at the metaphase are moving to the ends disappear. chromatids. Later plate. Although not in prometaphase, the of cell as their yet visible nuclear envelop will kinetochore in the micrograph, fragment. microtubles shorten. the mitotic spindle is staring to from. Figure 12.10 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Telophase. Daughter nuclei are forming. Meanwhile, cytokinesis has started: The cell plate, which will divided the cytoplasm in two, is growing toward the perimeter of the parent cell.
  • 27. Binary Fission • Prokaryotes (bacteria) – Reproduce by a type of cell division called binary fission Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 28. • In binary fission – The bacterial chromosome replicates – The two daughter chromosomes actively move apart Origin of replication Cell wall E. coli cell Chromosome replication 1 begins. Soon thereafter, one copy of the origin moves rapidly toward the other end of the cell. 2 Replication continues. One copy of the origin is now at each end of the cell. 3 Replication finishes. The plasma membrane grows inward, and new cell wall is deposited. Figure 12.11 4 Two daughter cells result. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Two copies of origin Origin Plasma Membrane Bacterial Chromosom e Origin
  • 29. The Evolution of Mitosis • Since prokaryotes preceded eukaryotes by billions of years – It is likely that mitosis evolved from bacterial cell division • Certain protists – Exhibit types of cell division that seem intermediate between binary fission and mitosis carried out by most eukaryotic cells Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 30. • A hypothetical sequence for the evolution of mitosis (a) Prokaryotes. During binary fission, the origins of the daughter chromosomes move to opposite ends of the cell. The mechanism is not fully understood, but proteins may anchor the daughter chromosomes to specific sites on the plasma membrane. (b) Dinoflagellates. In unicellular protists called dinoflagellates, the nuclear envelope remains intact during cell division, and the chromosomes attach to the nuclear envelope. Microtubules pass through the nucleus inside cytoplasmic tunnels, reinforcing the spatial orientation of the nucleus, which then divides in a fission process reminiscent of bacterial division. (c) Diatoms. In another group of unicellular protists, the diatoms, the nuclear envelope also remains intact during cell division. But in these organisms, the microtubules form a spindle within the nucleus. Microtubules separate the chromosomes, and the nucleus splits into two daughter nuclei. (d) Most eukaryotes. In most other eukaryotes, including plants and animals, the spindle forms outside the nucleus, and the nuclear envelope breaks down during mitosis. Microtubules separate the chromosomes, and the nuclear envelope then re-forms. Figure 12.12 A-D Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Bacterial chromosome Chromosomes Microtubules Intact nuclear envelope Kinetochore microtubules Intact nuclear envelope Kinetochore microtubules Centrosome Fragments of nuclear envelope
  • 31. • Concept 12.3: The 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 32. Evidence for Cytoplasmic Signals • Molecules present in the cytoplasm – Regulate progress through the cell cycle EXPERIMENTS In each experiment, cultured mammalian cells at two different phases of the cell cycle were induced to fuse. Experiment 2 Experiment 1 S G1 M S M G1 RESULTS S When a cell in the S phase was fused with a cell in G1, the G1 cell immediately entered the S phase—DNA was synthesized. M When a cell in the M phase was fused with a cell in G1, the G1 cell immediately began mitosis— a spindle formed and chromatin condensed, even though the chromosome had not been duplicated. CONCLUSION The results of fusing cells at two different phases of the cell cycle suggest that molecules present in the Figure 12.13 A, B cytoplasm of cells in the S or M phase control the progression of phases. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 33. 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 G1 checkpoint Control system G1 M G2 M checkpoint Figure 12.14 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings G2 checkpoint S
  • 34. • The clock has specific checkpoints – Where the cell cycle stops until a go-ahead signal is received G0 G1 checkpoint G1 (a) If a cell receives a go-ahead signal at the G1 checkpoint, the cell continues Figure 12.15 A, B on in the cell cycle. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings G1 (b) If a cell does not receive a go-ahead signal at the G1checkpoint, the cell exits the cell cycle and goes into G0, a nondividing state.
  • 35. 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) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 36. • The activity of cyclins and Cdks – Fluctuates during the cell cycle (a) Fluctuation of MPF activity and cyclin concentration during the cell cycle M G1 S G2 M G1 S G2 M MPF activity Cyclin Time (b) Molecular mechanisms that help regulate the cell cycle 1 Synthesis of cyclin begins in late S phase and continues through G2. Because cyclin is protected from degradation during this stage, it accumulates. G 4 During anaphase, the cyclin component of MPF is degraded, terminating the M phase. The cell enters the G1 phase. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 G Degraded Cyclin Cyclin is degraded Figure 12.16 A, B S Cdk M the cell favor degradation of cyclin, and the Cdk component of MPF is recycled. 1 5 During G1, conditions in G2 Cdk checkpoint MPF Cyclin 2 Accumulated cyclin molecules combine with recycled Cdk molecules, producing enough molecules of MPF to pass the G2 checkpoint and initiate the events of mitosis. 3 MPF promotes mitosis by phosphorylating various proteins. MPF‘s activity peaks during metaphase.
  • 37. Stop and Go Signs: Internal and External Signals at the Checkpoints • Both internal and external signals – Control the cell cycle checkpoints Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 38. • Growth factors – Stimulate other cells to divide EXPERIMENT Scalpels 1 A sample of connective tissue was cut up into small pieces. Petri plate 2 Enzymes were used to digest the extracellular matrix, resulting in a suspension of free fibroblast cells. 3 Cells were transferred to sterile culture vessels Figure 12.17 containing a basic growth medium consisting of glucose, amino acids, salts, and antibiotics (as a Without PDGF precaution against bacterial growth). PDGF was added to half the vessels. The culture vessels were incubated at 37°C. With PDGF Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 39. • In density-dependent inhibition – Crowded cells stop dividing • Most animal cells exhibit anchorage dependence – In which they must be attached to a substratum to divide (a) Normal mammalian cells. The availability of nutrients, growth factors, and a substratum for attachment limits cell density to a single layer. Cells anchor to dish surface and divide (anchorage dependence). When cells have formed a complete single layer, they stop dividing (density-dependent inhibition). If some cells are scraped away, the remaining cells divide to fill the gap and then stop (densitydependent inhibition). Figure 12.18 A Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 25 µm
  • 40. • Cancer cells – Exhibit neither density-dependent inhibition nor anchorage dependence Cancer cells do not exhibit anchorage dependence or density-dependent inhibition. (b) Cancer cells. Cancer cells usually continue to divide well beyond a single layer, forming a clump of overlapping cells. Figure 12.18 B 25 µm Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 41. Loss of Cell Cycle Controls in Cancer Cells • Cancer cells – Do not respond normally to the body’s control mechanisms – Form tumors Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
  • 42. • Malignant tumors invade surrounding tissues and can metastasize – Exporting cancer cells to other parts of the body where they may form secondary tumors Lymph vessel Tumor Blood vessel Glandular tissue 1 A tumor grows from a single cancer cell. Cancer cell 2 Cancer cells invade neighboring tissue. Figure 12.19 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 3 Cancer cells spread through lymph and blood vessels to other parts of the body. Metastatic Tumor 4 A small percentage of cancer cells may survive and establish a new tumor in another part of the body.