LECTURE PRESENTATIONS

For CAMPBELL BIOLOGY, NINTH EDITION

Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson

Chapter 12

The Cell Cycle

Lectures by
Erin Barley
Kathleen Fitzpatrick

© 2011 Pearson Education, Inc.

Overview: The Key Roles of Cell Division
• The ability of organisms to produce more of their
own kind best distinguishes living things from
nonliving matter
• The continuity of life is based on the reproduction
of cells, or cell division

© 2011 Pearson Education, Inc.

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
• Cell division is an integral part of the cell cycle,
the life of a cell from formation to its own division

© 2011 Pearson Education, Inc.

Figure 12.2
100 m (a) Reproduction

200 m
(b) Growth and
development

20 m
(c) Tissue renewal
Concept 12.1: Most cell division results in
genetically identical 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

© 2011 Pearson Education, Inc.

Cellular Organization of the 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

© 2011 Pearson Education, Inc.

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

© 2011 Pearson Education, Inc.

 Distribution of Chromosomes During
Eukaryotic Cell Division
• In preparation for 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

© 2011 Pearson Education, Inc.

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
Chromosomal
Chromosomes DNA molecules
1 Centromere

Chromosome
arm
Figure 12.5-2
Chromosomal
Chromosomes DNA molecules
1 Centromere

Chromosome
arm
Chromosome duplication
(including DNA replication)
and condensation
2

Sister
chromatids
Figure 12.5-3
Chromosomal
Chromosomes DNA molecules
1 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

© 2011 Pearson Education, Inc.

Concept 12.2: The mitotic phase alternates
with interphase in the cell cycle
• In 1882, the German anatomist Walther Flemming
developed dyes to observe chromosomes during
mitosis and cytokinesis

© 2011 Pearson Education, Inc.

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
– G1 phase (“first gap”)
– S phase (“synthesis”)
– G2 phase (“second gap”)
• The cell grows during all three phases, but
chromosomes are duplicated only during the S
phase

© 2011 Pearson Education, Inc.

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

© 2011 Pearson Education, Inc.

 https://www.youtube.com/watch?v=SEJuGFsNeBI

https://www.youtube.com/watch?v=mKWxeMMFTEU

BioFlix: Mitosis
© 2011 Pearson Education, Inc.
Figure 12.7

10 m
G2 of Interphase Prophase Prometaphase Metaphase Anaphase Telophase and Cytokinesis
Centrosomes Chromatin Fragments Nonkinetochore
(with centriole pairs) (duplicated) Early mitotic Aster of nuclear microtubules Metaphase Cleavage Nucleolus
spindle Centromere envelope plate furrow forming

Plasma
Nucleolus Nuclear membrane Chromosome, consisting Kinetochore Kinetochore Nuclear
envelope of two sister chromatids microtubule Spindle Centrosome at Daughter envelope
one spindle pole chromosomes forming
Figure 12.7a

G2 of Interphase Prophase Prometaphase

Centrosomes Fragments
(with centriole Chromatin Early mitotic Aster of nuclear Nonkinetochore
pairs) (duplicated) spindle envelope microtubules
Centromere

Plasma
Nucleolus membrane Kinetochore Kinetochore
Chromosome, consisting
Nuclear of two sister chromatids microtubule
envelope
Figure 12.7b

Metaphase Anaphase Telophase and Cytokinesis

Metaphase Cleavage Nucleolus

plate furrow forming

Nuclear
Spindle Centrosome at Daughter envelope
one spindle pole chromosomes forming
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
• The centrosome replicates during interphase,
forming two centrosomes that migrate to opposite
ends of the cell during prophase and
prometaphase

© 2011 Pearson Education, Inc.

 • An aster (a radial array of short microtubules)
extends from each centrosome
• The spindle includes the centrosomes, the spindle
microtubules, and the asters

© 2011 Pearson Education, Inc.

 • 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

Centrosome
Aster
Metaphase
Sister plate
chromatids (imaginary) Microtubules

Chromosomes
Kineto-
chores Centrosome
1 m

Overlapping
nonkinetochore
microtubules Kinetochore
microtubules

0.5 m
Figure 12.8a

Kinetochores

Kinetochore
microtubules

0.5 m
Figure 12.8b

Microtubules

Chromosomes

Centrosome

1 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

© 2011 Pearson Education, Inc.

Figure 12.9
EXPERIMENT
Kinetochore

Spindle
pole

Mark

RESULTS

CONCLUSION
Chromosome
movement
Microtubule Kinetochore

Motor protein Tubulin

subunits
Chromosome
Figure 12.9a
EXPERIMENT
Kinetochore

Spindle
pole

Mark

RESULTS
Figure 12.9b

CONCLUSION
Chromosome
movement
Microtubule Kinetochore

Motor protein Tubulin

subunits
Chromosome
 • 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
• Cytokinesis begins during anaphase or telophase
and the spindle eventually disassembles

© 2011 Pearson Education, Inc.

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

© 2011 Pearson Education, Inc.

Figure 12.10

(a) Cleavage of an animal cell (SEM) (b) Cell plate formation in a plant cell (TEM)

100 m
Cleavage furrow Vesicles Wall of parent cell
forming 1 m
cell plate Cell plate New cell wall

Contractile ring of Daughter cells

microfilaments
Daughter cells
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 12.12-1
Origin of Cell wall
replication Plasma membrane
E. coli cell
Bacterial chromosome
1 Chromosome Two copies
replication of origin
begins.
Figure 12.12-2
Origin of Cell wall
replication Plasma membrane
E. coli cell
Bacterial chromosome
1 Chromosome Two copies
replication of origin
begins.

2 Replication Origin Origin

continues.
Figure 12.12-3
Origin of Cell wall
replication Plasma membrane
E. coli cell
Bacterial chromosome
1 Chromosome Two copies
replication of origin
begins.

2 Replication Origin Origin

continues.

3 Replication
finishes.
Figure 12.12-4
Origin of Cell wall
replication Plasma membrane
E. coli cell
Bacterial chromosome
1 Chromosome Two copies
replication of origin
begins.

2 Replication Origin Origin

continues.

3 Replication
finishes.

4 Two daughter
cells result.
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
Bacterial
(a) Bacteria
chromosome

Chromosomes

Microtubules
(b) Dinoflagellates

Intact nuclear
envelope

Kinetochore
microtubule
(c) Diatoms and
some yeasts Intact nuclear
envelope

Kinetochore
microtubule
(d) Most eukaryotes
Fragments of
nuclear envelope
Figure 12.13a

Bacterial
chromosome

(a) Bacteria

Chromosomes

Microtubules

Intact nuclear
envelope
(b) Dinoflagellates
Figure 12.13b

Kinetochore
microtubule

Intact nuclear
envelope

(c) Diatoms and some yeasts

Kinetochore
microtubule

Fragments of
nuclear envelope
(d) Most eukaryotes
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 differences result from regulation at the
molecular level
• Cancer cells manage to escape the usual controls
on the cell cycle

© 2011 Pearson Education, Inc.

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

© 2011 Pearson Education, Inc.

Figure 12.14
EXPERIMENT
Experiment 1 Experiment 2

S G1 M G1

RESULTS

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

© 2011 Pearson Education, Inc.

Figure 12.15
G1 checkpoint

Control
system S
G1

M G2

M checkpoint
G2 checkpoint
 • For many cells, the G1 checkpoint seems to be the
most important
• 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

© 2011 Pearson Education, Inc.

Figure 12.16

G0
G1 checkpoint

G1 G1

(a) Cell receives a go-ahead (b) Cell does not receive a

signal. 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 G2
checkpoint into the M phase
© 2011 Pearson Education, Inc.
Figure 12.17a

M G 1 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
Figure 12.17b

Cdk

Degraded
cyclin G2 Cdk
checkpoint
Cyclin is
degraded
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
• 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

© 2011 Pearson Education, Inc.

Figure 12.18

1 A sample of human Scalpels

connective tissue is
cut up into small
pieces.
Petri
dish
2 Enzymes digest
the extracellular
matrix, resulting in
a suspension of
free fibroblasts.
4 PDGF is added 10 m
3 Cells are transferred to to half the
culture vessels. vessels.

Without PDGF With PDGF

Figure 12.18a

10 m
 • A clear 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
• 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
Figure 12.19a

20 m
Figure 12.19b

20 m
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

© 2011 Pearson Education, Inc.

 • 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

Lymph
vessel
Tumor
Blood
vessel

Glandular Cancer
tissue cell
Metastatic
tumor
1 A tumor grows 2 Cancer 3 Cancer cells spread 4 Cancer cells
from a single cells invade through lymph and may survive
cancer cell. neighboring blood vessels to and establish
tissue. other parts of the a new tumor
body. in another part
of the body.
 • Recent advances in understanding the cell
cycle and cell cycle signaling have led to
advances in cancer treatment

© 2011 Pearson Education, Inc.

Figure 12.21
Figure 12.UN01
P

G1 S
Cytokinesis
Mitosis G2

MITOTIC (M) PHASE

Prophase
Telophase and
Cytokinesis

Prometaphase
Anaphase
Metaphase
MEIOSIS
1. Prophase I
1) Leptotene: inti mulai membesar,
kromosom memanjang, benang
kromatin belum teratur
2) Zygotene: kromosom homolog 2. Metaphase I
mengadakan synapsis, kromosom
memendek atau mengkerut
3) Pachytene: synapsis mjd
sempurna, pendek, tebal, tiap
kromosom membelah menjadi 2 shg
terbentuk tetrad
4) Diplotene: tetrad mulai membelah,
masing-masing belahan ada 2
kromatid tp msh tdp perlekatan yg
disbt chiasma
5) Diakenesis: pemisahan tetrad 3. Anaphase I
menjadi diad hampir sempurna dan 4. Telophase I
kromosom mengatur diri di bidang
ekuator sel
Interphase I

• Similar to mitosis interphase.

• Chromosomes replicate (S phase).

• Each duplicated chromosome consist of two

identical sister chromatids attached at their
centromeres.

• Centriole pairs also replicate.

 Interphase I

• Nucleus and nucleolus visible.

chromatin nuclear
membrane

cell membrane

nucleolus
Meiosis I (four phases)

• Cell division that reduces the chromosome

number by one-half.

• four phases:
a. prophase I
b. metaphase I
c. anaphase I
d. telophase I
 Prophase I

• Longest and most complex phase.

• 90% of the meiotic process is spent in Prophase I
• Chromosomes condense.
• Synapsis occurs: homologous chromosomes come
together to form a tetrad.
• Tetrad is two chromosomes or four chromatids
(sister and nonsister chromatids).
 Prophase I - Synapsis
Homologous chromosomes

sister chromatids sister chromatids

Tetrad
During Prophase I
“Crossing Over” occurs.
Crossing Over is one of the Two major
occurrences of Meiosis
(The other is Non-disjunction)
• During Crossing over segments of nonsister
chromatids break and reattach to the other
chromatid. The Chiasmata (chiasma) are the
sites of crossing over.
Crossing Over
creates variation (diversity) in the offspring’s traits.
nonsister chromatids Tetrad

chiasmata: site variation

of crossing over
Prophase I

spindle fiber centrioles

aster
fibers
 Metaphase I
• Shortest phase
• Tetrads align on the metaphase plate.
• INDEPENDENT ASSORTMENT OCCURS:
1. Orientation of homologous pair to poles is random.
2. Variation
3. Formula: 2n
Example: 2n = 4
then n = 2
thus 22 = 4 combinations
Metaphase I

OR

metaphase plate metaphase plate

Anaphase I

• Homologous chromosomes separate and

move towards the poles.

• Sister chromatids remain attached at their

centromeres.
Anaphase I
Telophase I

• Each pole now has haploid set of

chromosomes.

• Cytokinesis occurs and two haploid daughter

cells are formed.
Telophase I
Meiosis II

• No interphase II
(or very short - no more DNA replication)

• Remember: Meiosis II is similar to mitosis

Prophase II

• same as prophase in mitosis

Metaphase II

• same as metaphase in mitosis

metaphase plate metaphase plate

Anaphase II

• same as anaphase in mitosis

• sister chromatids separate
 Telophase II

• Same as telophase in mitosis.

• Nuclei form.
• Cytokinesis occurs.

• Remember: four haploid daughter cells

produced.

gametes = sperm or egg

Telophase II
Figure 9.17 Mitosis and Meiosis: A Comparison

Prophase

No synapsis of
MITOSIS homologous
chromosomes

Homologous
chromosome
pairs Synapsis and
MEIOSIS crossing over
of homologs

Crossover

Prophase I
Figure 9.17 Mitosis and Meiosis: A Comparison

Metaphase

Individual
MITOSIS chromosomes
align at the
equatorial plate.

Homologous
MEIOSIS pairs align at the
equatorial plate.

Metaphase I
Figure 9.17 Mitosis and Meiosis: A Comparison

Anaphase

Centromeres
MITOSIS separate. Sister
chromatids
separate during
anaphase, becoming
daughter
chromosomes.

Centromeres
MEIOSIS do not separate;
sister chromatids
remain together
during anaphase;
homologs separate;
DNA does not
Anaphase I replicate before
subsequent
prophase.
Figure 9.17 Mitosis and Meiosis: A Comparison

Two daughter cells (each 2n)

MITOSIS

MEIOSIS

Telophase I
Figure 9.17 Mitosis and Meiosis: A Comparison

No further division

MITOSIS

MEIOSIS

Metaphase II
Figure 9.17 Mitosis and Meiosis: A Comparison

No further division

MITOSIS

Chromatids
MEIOSIS separate.

Four daughter cells (each n)