Chapter 12. The Cell Cycle

Содержание

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Overview: The Key Roles of Cell Division The ability of organisms

Overview: The Key Roles of Cell Division

The ability of organisms to

reproduce best distinguishes living things from nonliving matter
The continuity of life is based on the reproduction of cells, or cell division

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Fig. 12-1

Fig. 12-1

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In unicellular organisms, division of one cell reproduces the entire organism

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

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Fig. 12-2 100 µm 200 µm 20 µm (a) Reproduction (b)

Fig. 12-2

100 µm

200 µm

20 µm

(a) Reproduction

(b) Growth and
development

(c) Tissue renewal

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Fig. 12-2a 100 µm (a) Reproduction

Fig. 12-2a

100 µm

(a) Reproduction

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Fig. 12-2b 200 µm (b) Growth and development

Fig. 12-2b

200 µm

(b) Growth and development

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Fig. 12-2c 20 µm (c) Tissue renewal

Fig. 12-2c

20 µm

(c) Tissue renewal

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Concept 12.1: Cell division results in genetically identical daughter cells Most

Concept 12.1: Cell division results in genetically identical daughter cells

Most cell

division results in daughter cells with identical genetic information, DNA
A special type of division produces nonidentical daughter cells (gametes, or sperm and egg cells)

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Cellular Organization of the Genetic Material All the DNA in a

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

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Fig. 12-3 20 µm

Fig. 12-3

20 µm

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Every eukaryotic species has a characteristic number of chromosomes in each

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
Eukaryotic chromosomes consist of chromatin, a complex of DNA and protein that condenses during cell division

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Distribution of Chromosomes During Eukaryotic Cell Division In preparation for cell

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, 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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Fig. 12-4 0.5 µm Chromosomes Chromosome duplication (including DNA synthesis) Chromo-

Fig. 12-4

0.5 µm

Chromosomes

Chromosome
duplication
(including DNA
synthesis)

Chromo-
some arm

Centromere

Sister
chromatids

DNA molecules

Separation of
sister chromatids

Centromere

Sister chromatids

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Eukaryotic cell division consists of: Mitosis, the division of the nucleus

Eukaryotic cell division consists of:
Mitosis, the division of 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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Concept 12.2: The mitotic phase alternates with interphase in the cell

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

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Phases of the Cell Cycle The cell cycle consists of Mitotic

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

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

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Fig. 12-5 S (DNA synthesis) MITOTIC (M) PHASE Mitosis Cytokinesis G1 G2 INTERPHASE

Fig. 12-5

S
(DNA synthesis)

MITOTIC
(M) PHASE

Mitosis

Cytokinesis

G1

G2

INTERPHASE

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Mitosis is conventionally divided into five phases: Prophase Prometaphase Metaphase Anaphase

Mitosis is conventionally divided into five phases:
Prophase
Prometaphase
Metaphase
Anaphase
Telophase
Cytokinesis is well underway by

late telophase

BioFlix: Mitosis

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Fig. 12-6 G2 of Interphase Centrosomes (with centriole pairs) Chromatin (duplicated)

Fig. 12-6

G2 of Interphase

Centrosomes
(with centriole
pairs)

Chromatin
(duplicated)

Nucleolus

Nuclear
envelope

Plasma
membrane

Early mitotic
spindle

Aster

Centromere

Chromosome, consisting
of two sister chromatids

Prophase

Prometaphase

Fragments
of

nuclear
envelope

Nonkinetochore
microtubules

Kinetochore

Kinetochore
microtubule

Metaphase

Metaphase
plate

Spindle

Centrosome at
one spindle pole

Anaphase

Daughter
chromosomes

Telophase and Cytokinesis

Cleavage
furrow

Nucleolus
forming

Nuclear
envelope
forming

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Prophase Fig. 12-6a Prometaphase G2 of Interphase

Prophase

Fig. 12-6a

Prometaphase

G2 of Interphase

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Fig. 12-6b Prometaphase Prophase G2 of Interphase Nonkinetochore microtubules Fragments of

Fig. 12-6b

Prometaphase

Prophase

G2 of Interphase

Nonkinetochore
microtubules

Fragments
of nuclear
envelope

Aster

Centromere

Early mitotic
spindle

Chromatin
(duplicated)

Centrosomes
(with centriole
pairs)

Nucleolus

Nuclear
envelope

Plasma
membrane

Chromosome, consisting
of two sister chromatids

Kinetochore

Kinetochore
microtubule

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Fig. 12-6c Metaphase Anaphase Telophase and Cytokinesis

Fig. 12-6c

Metaphase

Anaphase

Telophase and Cytokinesis

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Fig. 12-6d Metaphase Anaphase Telophase and Cytokinesis Cleavage furrow Nucleolus forming

Fig. 12-6d

Metaphase

Anaphase

Telophase and Cytokinesis

Cleavage
furrow

Nucleolus
forming

Metaphase
plate

Centrosome at
one spindle pole

Spindle

Daughter
chromosomes

Nuclear
envelope
forming

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The Mitotic Spindle: A Closer Look The mitotic spindle is an

The Mitotic Spindle: A Closer Look

The mitotic spindle is an apparatus

of microtubules that controls chromosome movement during mitosis
During prophase, assembly of spindle microtubules begins in the centrosome, the microtubule organizing center
The centrosome replicates, forming two centrosomes that migrate to opposite ends of the cell, as spindle microtubules grow out from them

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An aster (a radial array of short microtubules) extends from each

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

During prometaphase, some spindle microtubules attach to the kinetochores of chromosomes

and begin to move the chromosomes
At metaphase, the chromosomes are all lined up at the metaphase plate, the midway point between the spindle’s two poles

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Fig. 12-7 Microtubules Chromosomes Sister chromatids Aster Metaphase plate Centrosome Kineto-

Fig. 12-7

Microtubules

Chromosomes

Sister
chromatids

Aster

Metaphase
plate

Centrosome

Kineto-
chores

Kinetochore
microtubules

Overlapping
nonkinetochore
microtubules

Centrosome

1 µm

0.5 µm

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In anaphase, sister chromatids separate and move along the kinetochore microtubules

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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Fig. 12-8 EXPERIMENT Kinetochore RESULTS CONCLUSION Spindle pole Mark Chromosome movement

Fig. 12-8

EXPERIMENT

Kinetochore

RESULTS

CONCLUSION

Spindle
pole

Mark

Chromosome
movement

Kinetochore

Microtubule

Motor
protein

Chromosome

Tubulin
subunits

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Fig. 12-8a Kinetochore Spindle pole Mark EXPERIMENT RESULTS

Fig. 12-8a

Kinetochore

Spindle
pole

Mark

EXPERIMENT

RESULTS

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Fig. 12-8b Kinetochore Microtubule Tubulin Subunits Chromosome Chromosome movement Motor protein CONCLUSION

Fig. 12-8b

Kinetochore

Microtubule

Tubulin
Subunits

Chromosome

Chromosome
movement

Motor
protein

CONCLUSION

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Nonkinetochore microtubules from opposite poles overlap and push against each other,

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

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Cytokinesis: A Closer Look In animal cells, cytokinesis occurs by a

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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Video: Sea Urchin (Time Lapse) Video: Animal Mitosis Copyright © 2008

Video: Sea Urchin (Time Lapse)

Video: Animal Mitosis

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Inc., publishing as Pearson Benjamin Cummings
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Fig. 12-9 Cleavage furrow 100 µm Contractile ring of microfilaments Daughter

Fig. 12-9

Cleavage furrow

100 µm

Contractile ring of
microfilaments

Daughter cells

(a) Cleavage of an animal

cell (SEM)

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

Vesicles
forming
cell plate

Wall of
parent cell

Cell plate

Daughter cells

New cell wall

1 µm

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Cleavage furrow Fig. 12-9a 100 µm Daughter cells (a) Cleavage of

Cleavage furrow

Fig. 12-9a

100 µm

Daughter cells

(a) Cleavage of an animal cell (SEM)

Contractile

ring of
microfilaments
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Fig. 12-9b Daughter cells (b) Cell plate formation in a plant

Fig. 12-9b

Daughter cells

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

Vesicles
forming
cell

plate

Wall of
parent cell

New cell wall

Cell plate

1 µm

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Fig. 12-10 Chromatin condensing Metaphase Anaphase Telophase Prometaphase Nucleus Prophase 1

Fig. 12-10

Chromatin
condensing

Metaphase

Anaphase

Telophase

Prometaphase

Nucleus

Prophase

1

2

3

5

4

Nucleolus

Chromosomes

Cell plate

10 µm

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Fig. 12-10a Nucleus Prophase 1 Nucleolus Chromatin condensing

Fig. 12-10a

Nucleus

Prophase

1

Nucleolus

Chromatin
condensing

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Fig. 12-10b Prometaphase 2 Chromosomes

Fig. 12-10b

Prometaphase

2

Chromosomes

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Fig. 12-10c Metaphase 3

Fig. 12-10c

Metaphase

3

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Fig. 12-10d Anaphase 4

Fig. 12-10d

Anaphase

4

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Fig. 12-10e Telophase 5 Cell plate 10 µm

Fig. 12-10e

Telophase

5

Cell plate

10 µm

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Binary Fission Prokaryotes (bacteria and archaea) reproduce by a type of

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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Fig. 12-11-1 Origin of replication Two copies of origin E. coli

Fig. 12-11-1

Origin of
replication

Two copies
of origin

E. coli cell

Bacterial
chromosome

Plasma
membrane

Cell wall

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Fig. 12-11-2 Origin of replication Two copies of origin E. coli

Fig. 12-11-2

Origin of
replication

Two copies
of origin

E. coli cell

Bacterial
chromosome

Plasma
membrane

Cell wall

Origin

Origin

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Fig. 12-11-3 Origin of replication Two copies of origin E. coli

Fig. 12-11-3

Origin of
replication

Two copies
of origin

E. coli cell

Bacterial
chromosome

Plasma
membrane

Cell wall

Origin

Origin

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Fig. 12-11-4 Origin of replication Two copies of origin E. coli

Fig. 12-11-4

Origin of
replication

Two copies
of origin

E. coli cell

Bacterial
chromosome

Plasma
membrane

Cell wall

Origin

Origin

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The Evolution of Mitosis Since prokaryotes evolved before eukaryotes, mitosis probably

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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Fig. 12-12 (a) Bacteria Bacterial chromosome Chromosomes Microtubules Intact nuclear envelope

Fig. 12-12

(a) Bacteria

Bacterial
chromosome

Chromosomes

Microtubules

Intact nuclear
envelope

(b) Dinoflagellates

Kinetochore
microtubule

Intact nuclear
envelope

(c) Diatoms and yeasts

Kinetochore
microtubule

Fragments of
nuclear envelope

(d)

Most eukaryotes
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Fig. 12-12ab Bacterial chromosome Chromosomes Microtubules (a) Bacteria (b) Dinoflagellates Intact nuclear envelope

Fig. 12-12ab

Bacterial
chromosome

Chromosomes

Microtubules

(a) Bacteria

(b) Dinoflagellates

Intact nuclear
envelope

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Fig. 12-12cd Kinetochore microtubule (c) Diatoms and yeasts Kinetochore microtubule (d)

Fig. 12-12cd

Kinetochore
microtubule

(c) Diatoms and yeasts

Kinetochore
microtubule

(d) Most eukaryotes

Fragments of
nuclear envelope

Intact nuclear
envelope

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Concept 12.3: The eukaryotic cell cycle is regulated by a molecular

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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Evidence for Cytoplasmic Signals The cell cycle appears to be driven

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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Fig. 12-13 Experiment 1 Experiment 2 EXPERIMENT RESULTS S G1 M

Fig. 12-13

Experiment 1

Experiment 2

EXPERIMENT

RESULTS

S

G1

M

G1

M

M

S

S

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.

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The Cell Cycle Control System The sequential events of the cell

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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Fig. 12-14 S G1 M checkpoint G2 M Control system G1 checkpoint G2 checkpoint

Fig. 12-14

S

G1

M checkpoint

G2

M

Control
system

G1 checkpoint

G2 checkpoint

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For many cells, the G1 checkpoint seems to be the most

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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Fig. 12-15 G1 G0 G1 checkpoint Cell receives a go-ahead signal

Fig. 12-15

G1

G0

G1 checkpoint

Cell receives a go-ahead
signal

G1

(b) Cell does not receive

a
go-ahead signal
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The Cell Cycle Clock: Cyclins and Cyclin-Dependent Kinases Two types of

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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Fig. 12-16 Protein kinase activity (– ) % of dividing cells

Fig. 12-16

Protein kinase activity (– )

% of dividing cells (– )

Time

(min)

300

200

400

100

0

1

2

3

4

5

30

500

0

20

10

RESULTS

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Fig. 12-17 M G1 S G2 M G1 S G2 M

Fig. 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

Degraded
cyclin

Cdk

G1

S

G2

M

Cdk

G2

checkpoint

Cyclin is
degraded

Cyclin

MPF

(b) Molecular mechanisms that help regulate the cell cycle

Cyclin accumulation

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Fig. 12-17a Time (a) Fluctuation of MPF activity and cyclin concentration

Fig. 12-17a

Time

(a) Fluctuation of MPF activity and cyclin concentration during
the

cell cycle

Cyclin
concentration

MPF activity

M

M

M

S

S

G1

G1

G1

G2

G2

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Fig. 12-17b Cyclin is degraded Cdk MPF Cdk M S G1

Fig. 12-17b

Cyclin is
degraded

Cdk

MPF

Cdk

M

S

G1

G2
checkpoint

Degraded
cyclin

Cyclin

(b) Molecular mechanisms that help regulate the cell cycle

G2

Cyclin

accumulation
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Stop and Go Signs: Internal and External Signals at the Checkpoints

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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Fig. 12-18 Petri plate Scalpels Cultured fibroblasts Without PDGF cells fail

Fig. 12-18

Petri
plate

Scalpels

Cultured fibroblasts

Without PDGF
cells fail to divide

With PDGF
cells prolifer-
ate

10 µm

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Another example of external signals is density-dependent inhibition, in which crowded

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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Fig. 12-19 Anchorage dependence Density-dependent inhibition Density-dependent inhibition (a) Normal mammalian

Fig. 12-19

Anchorage dependence

Density-dependent inhibition

Density-dependent inhibition

(a) Normal mammalian cells

(b) Cancer cells

25 µm

25

µm
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Cancer cells exhibit neither density-dependent inhibition nor anchorage dependence Copyright ©

Cancer cells exhibit neither density-dependent inhibition nor anchorage dependence

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Loss of Cell Cycle Controls in Cancer Cells Cancer cells do

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

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

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Fig. 12-20 Tumor A tumor grows from a single cancer cell.

Fig. 12-20

Tumor

A tumor grows
from a single
cancer cell.

Glandular
tissue

Lymph
vessel

Blood
vessel

Metastatic
tumor

Cancer
cell

Cancer cells
invade neigh-
boring tissue.

Cancer cells

spread
to other parts of
the body.

Cancer cells may
survive and
establish a new
tumor in another
part of the body.

1

2

3

4

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Fig. 12-UN1 Telophase and Cytokinesis Anaphase Metaphase Prometaphase Prophase MITOTIC (M)

Fig. 12-UN1

Telophase and
Cytokinesis

Anaphase

Metaphase

Prometaphase

Prophase

MITOTIC (M) PHASE

Cytokinesis

Mitosis

S

G1

G2

INTERPHASE

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Fig. 12-UN2

Fig. 12-UN2

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Fig. 12-UN3

Fig. 12-UN3

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Fig. 12-UN4

Fig. 12-UN4

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Fig. 12-UN5

Fig. 12-UN5

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Fig. 12-UN6

Fig. 12-UN6

Слайд 81

You should now be able to: Describe the structural organization of

You should now be able to:

Describe the structural organization of the

prokaryotic genome and the eukaryotic genome
List the phases of the cell cycle; describe the sequence of events during each phase
List the phases of mitosis and describe the events characteristic of each phase
Draw or describe the mitotic spindle, including centrosomes, kinetochore microtubules, nonkinetochore microtubules, and asters

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