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CAMPBELL BIOLOGY IN FOCUS

Urry Cain Wasserman Minorsky Jackson Reece

9
The Cell Cycle

Lecture Presentations by
Kathleen Fitzpatrick and Nicole Tunbridge
Edited by Rena Quinlan, Ph.D.
2014 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
Figure 9.1. This series of fluorescence micrographs follows an
animal cells chromosomes, from lower left to lower right, as one
cell divides into two.
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In unicellular organisms,
division of one cell
reproduces the entire
organism

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

(a) Reproduction

Cell division enables


multicellular eukaryotes
to develop from a single
cell and, once fully
grown, to renew, repair,
or replace cells as
needed
Cell division is an
integral part of the cell
cycle, the life of a cell
from formation to its own
division
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200 m
(b) Growth and
development. This
micrograph shows a
sand dollar embryo
shortly after the fertilized
egg divided, forming two
cells (LM).

(b) Growth and development


(c) Tissue renewal. These
dividing bone marrow cells
will give rise to new blood
cells (LM).

20 m

(c) Tissue renewal

Fig. 9.2. The functions of cell


division.

Concept 9.1: Most cell division results in


genetically identical daughter cells
Most cell division results in the distribution of
identical genetic materialDNAto two daughter
cells
DNA is passed from one generation of cells to the
next with remarkable fidelity

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Cellular Organization of the Genetic Material


All the DNA in a cell
constitutes the cells
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 (Fig. 9.3)
Eukaryotic chromosomes
consist of chromatin, a
complex of DNA and
protein

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20 m
Fig. 9.3 Eukaryotic chromosomes. Chromosomes (stained purple)
are visible within the nucleus of this cell from an African blood lily.
The thinner red threads in the surrounding cytoplasm are the
cytoskeleton. The cell is preparing to divide (LM).

Every eukaryotic species has a characteristic


number of chromosomes in each cell nucleus
Somatic cells (non-reproductive cells) have two sets
of chromosomes DIPLOID (2n)
Two sets total: one set from each parent

Gametes (reproductive cells: sperm and eggs) have


one set of chromosomes HAPLOID (n)

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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 identical copies of
the original chromosome
The centromere is where the two chromatids are most closely attached

Sister
chromatids

Centromere

0.5 m

Fig. 9.4. A highly condensed, duplicated human chromosome (SEM).


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Chromosomal
DNA molecules

Chromosomes
1

During cell
division, the
two sister
chromatids
of each
duplicated
chromosome
separate and
move into two
nuclei

Centromere

Chromosome
arm
Chromosome duplication
2

Sister
chromatids
Separation of sister
chromatids
3

Fig. 9.5. Chromosome duplication and distribution during cell division.


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Eukaryotic cell division consists of


Mitosis, the division of the genetic material in the
nucleus
Cytokinesis, the division of the cytoplasm

Gametes (sex cells; egg and sperm) 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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Concept 9.2: The mitotic phase alternates with


interphase in the cell cycle

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Phases of the Cell Cycle


The cell cycle consists of:
Mitotic (M) phase,
including mitosis and
cytokinesis
Interphase, including cell
growth and copying of
chromosomes in
preparation for cell
division

G1

Interphase (about 90% of the


cell cycle) can be divided into
subphases:

s
si
e
n
ki
o
t
Cy

ito

si
s

G1 phase (first gap)

S phase (DNA
synthesis)
G2 phase (second gap)
During all three phases of
Interphase the cell grows by
producing proteins and
organelles (ie., mitochondria,
ER), but chromosomes are
duplicated only during the S
phase
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Figure 9.6

S
(DNA synthesis)

G2

Mitosis is divided into five main phases

10 m

2nd Stage: Pro-metaphase = nuclear


envelope fragments and spindle
microtubules attach to the kinetochores
of the chromosomes
- Kinetochore: a structure of proteins
attached to the centromere that links each
sister chromatid to the mitotic spindle

10 m

1st Stage: Prophase = chromatin condenses


into discrete chromosomes, mitotic
spindle begins to form

Prophase

Prometaphase

10 m

4th Stage: Anaphase = chromatids of each


chromosome have separated and
daughter chromosomes are moving to the
opposite poles of the cell.

10 m

3rd Stage: Metaphase = Spindle is complete


and the chromosomes, attached to
microtubules at their kinetochores, align
at the metaphase plate.

Anaphase

Metaphase

10 m

5th Stage: Telophase = daughter nuclei are


forming and cytokinesis has begun
- Cytokinesis: division of cytoplasm to
form two separate daughter cells
immediately after mitosis
Telophase and
Cytokinesis

10 m

Figure 9.7a

G2 of Interphase
Centrosomes
(with centriole
pairs)

Chromosomes Early mitotic


Centromere
(duplicated,
spindle
Aster
uncondensed)

Plasma
Nucleolus Nuclear membrane
envelope
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Prophase

Two sister chromatids of


one chromosome

Prometaphase
Fragments
of nuclear
envelope

Kinetochore

Nonkinetochore
microtubules

Kinetochore
microtubule

10 m

Figure 9.7b

Metaphase

Anaphase

Metaphase
plate

Spindle

Centrosome at
one spindle pole

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Telophase and
Cytokinesis
Cleavage
furrow

Daughter
chromosomes

Nuclear
envelope
forming

Nucleolus
forming

The Mitotic Spindle: a stucture made of microtubules and associated


proteins that controls chromosome movement during mitosis

In animal cells, assembly of


spindle microtubules begins
in the centrosome, the
microtubule organizing
center

The spindle includes the


centrosomes, the spindle
microtubules, and the asters

During prometaphase, some


spindle microtubules attach
to the kinetochores (protein
complexes attached to
centromeres of each sister
chromatid) of chromosomes
and begin to move the
chromosomes

At metaphase, the
centromeres of all the
chromosomes are at the
metaphase plate, an
imaginary structure midway
between the spindles two
poles

radial array of short microtubules at opposite poles


during prophase and prometaphase
Aster

Sister
chromatids

Centrosome
Metaphase plate (imaginary)

Kinetochores
Microtubules Chromosomes

Overlapping
nonkinetochore
microtubules

Kinetochore
microtubules
1 m
0.5 m

Centrosome

Fig. 9.8. The mitotic spindle at metaphase. The kinetochores of each chromosomes two
sister chromatids face in opposite directions. Here, each kinetochore is attached to a
cluster of kinetochore microtubules extending from the nearest centrosome. (TEMs)

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
(a) Cleavage of an animal cell (SEM)

Cleavage furrow

Contractile ring of
microfilaments

100 m

(b) Cell plate formation in a plant cell (TEM)

Vesicles
forming
cell plate

Wall of parent cell

1 m

Cell plate

New cell wall

Daughter cells
Daughter cells

Figure 9.10
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Binary Fission in Bacteria


Prokaryotes (bacteria
and archaea)
reproduce by a type of
cell division called
binary fission
In E. coli, the single
chromosome
replicates, beginning at
the origin of
replication

Origin of
replication
E. coli cell
1 Chromosome
replication
begins.

2 One copy of the


origin is now at
each end of the
cell.

Cell wall
Plasma
membrane
Bacterial
chromosome

Two copies
of origin

Origin

Origin

Where it starts

The two daughter


chromosomes actively
move apart while the
cell elongates
The plasma membrane
pinches inward,
dividing the cell into two
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3 Replication
finishes.

4 Two daughter
cells result.
Figure 9.12. Bacterial cell division by binary fission. The
bacterium E. coli, shown here, has a single, circular
chromosome.

Concept 9.3: 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

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Checkpoints of the Cell Cycle Control System: the


sequential events of the cell cycle are directed by a cell
cycle control system
Both internal and
external signals control
cell cycle checkpoints
via signal transduction
pathways; key
regulatory proteins
are Kinases (activate
or inactivate proteins
by phosphorylating
them), and Cyclins
(proteins that regulate
the activity of cyclindependent kinases
(CDKs) .
The clock has specific
checkpoints where
the cell cycle stops
until a go-ahead signal
is received
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G1 checkpoint

Control
system

G1
M

M checkpoint

G2

G2 checkpoint
Figure 9.15

The G1 checkpoint: the most important checkpoint of


the Cell Cycle

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
Most human body cells are in the G0 phase (ie., mature nerve cells and muscle cells never divide), but
other cells, such as liver cells can re-enter the cell cycle if external cues (eg., growth factor proteins)
are released during injury.

G1 checkpoint

G0

G1
Without go-ahead signal,
cell enters G0.

G1
With go-ahead signal,
cell continues cell cycle.

(a) G1 checkpoint
Figure 9.16. The G1 checkpoint of the Cell Cycle

The M checkpoint; another important checkpoint of the


Cell Cycle: a cell in mitosis receives a stop signal when any
of its chromosomes are not attached to spindle fibers
An internal signal occurs
at the M phase
G
G
checkpoint
In this case, anaphase
M G
M G
does not begin if any
kinetochores remain
unattached to spindle M checkpoint
G
microtubules
checkpoint
Anaphase
Prometaphase
Attachment of all of the
Metaphase
A cell in mitosis receives a stop
kinetochores activates a
With full chromosome attachment
signal when any of its chromosomes
regulatory complex,
to spindle fibers from both poles,
are not attached to spindle fibers.
a go-ahead signal allows the cell to
which then activates the
proceed into anaphase.
enzyme separase
(b) M checkpoint
Separase allows sister
Fig. 9.16b. The M checkpoint of the Cell Cycle
chromatids to separate,
triggering the onset of
anaphase
1

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Density-dependent inhibition and anchorage dependence


of cell division
An example of an
external signal 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 densitydependent inhibition
nor anchorage
dependence

Anchorage dependence: cells


require a surface for division

Density-dependent inhibition:
when cells form a single layer
they stop dividing
Density-dependent inhibition:
if some cells are removed the remaining cells divide
to fill the gap and then stop once they contact each other

20 m

a) Normal mammalian cells.


Contact with neighboring cells and the
availability of nutrients, growth factors,
and a substratum for attachment limit
cell density to a single layer

20 m

(b) Cancer cells.


Cancer cells usually continue to
divide well beyond a single layer,
forming a clump of overlapping cells

Fig. 9.18. Density-dependent inhibition and anchorage dependence of cell division.


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Loss of Cell Cycle Controls in Cancer Cells


Cancer cells do not respond to signals that normally
regulate the cell cycle
Cancer cells may not need growth factors to grow
and divide
They may make their own growth factor
They may convey a growth factors signal without the
presence of the growth factor
They may have an abnormal cell cycle control system
Recent advances in understanding the cell cycle and cell cycle
signaling have led to advances in cancer treatment
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Fig. 9.19. The growth and metastasis of a malignant breast tumor.


A normal cell is converted to a cancerous cell by a process called transformation

Cancer cells 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
5 m

Breast cancer cell


(colorized SEM)

Lymph
vessel

Tumor

Blood
vessel

Glandular
tissue
1
Fig. 9.19

A tumor grows
from a single
cancer cell.

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Metastatic
tumor

Cancer cells
invade
neighboring tissue.

Cancer
cell
Cancer cells spread
4
through lymph and
blood vessels to
other parts of the body.

A small percentage
of cancer cells may
metastasize to another
part of the body.