7

The Cell Cycle and Cell

Division

Chapter 7 The Cell Cycle and Cell Division

Key Concepts
7.1 Different Life Cycles Use Different
Modes of Cell Reproduction
7.2 Both Binary Fission and Mitosis
Produce Genetically Identical Cells
7.3 Cell Reproduction Is Under Precise
Control

Chapter 7 The Cell Cycle and Cell Division

Key Concepts
7.4 Meiosis Halves the Nuclear
Chromosome Content and Generates
Diversity
7.5 Programmed Cell Death Is a
Necessary Process in Living Organisms

Chapter 7 Opening Question

How does infection with HPV result in

uncontrolled cell reproduction?

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

The lifespan of an organism is linked to cell

reproductionusually called cell division.

Organisms have two basic strategies for

reproducing themselves:
Asexual reproduction
Sexual reproduction
Cell division is also important in growth and
repair of tissues.

Figure 7.1 The Importance of Cell Division (Part 1)

Figure 7.1 The Importance of Cell Division (Part 2)

Figure 7.1 The Importance of Cell Division (Part 3)

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

In asexual reproduction the offspring are

clonesgenetically identical to the parent.
Any genetic variations are due to mutations.
A unicellular prokaryote may reproduce itself by
binary fission.
Single-cell eukaryotes can reproduce by mitosis.

Other eukaryotes are also able to reproduce

through asexual or sexual means.

Figure 7.2 Asexual Reproduction on a Large Scale

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

Sexual reproduction requires gametestwo

parents each contribute one gamete to an
offspring.
Gametes form by meiosisa process of cell
division.
Gametesand offspringdiffer genetically from
each other and from the parents.

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

DNA in eukaryotic cells is organized into

chromosomes.
A chromosome consists of a single molecule of
DNA and proteins.

Somatic cellsbody cells not specialized for

reproduction
Each somatic cell contains two sets of
chromosomes (homologs) that occur in
homologous pairs.

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

Gametes contain only one set of

chromosomesone homolog from each pair.
Haploid cellNumber of chromosomes = n
FertilizationTwo haploid gametes (female egg
and male sperm) fuse to form a zygote.
Chromosome number in zygote = 2n and cells
are diploid.

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

All kinds of sexual life cycles involve meiosis:

Haplontic life cyclein protists, fungi, and some

algaezygote is only diploid stage
After zygote forms it undergoes meiosis to form
haploid spores, which germinate to form a new
organism.
Organism is haploid, and produces gametes by
mitosiscells fuse to form diploid zygote.

Figure 7.3 All Sexual Life Cycles Involve Fertilization and Meiosis (Part 1)

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

Alternation of generationsmost plants, some

protists; meiosis gives rise to haploid spores
Spores divide by mitosis to form the haploid
generation (gametophyte).

Gametophyte forms gametes by mitosis.

Gametes then fuse to form diploid zygote
(sporophyte), which in turn produces haploid
spores by meiosis.

Figure 7.3 All Sexual Life Cycles Involve Fertilization and Meiosis (Part 2)

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

Diplontic life cycleanimals and some plants;

gametes are the only haploid stage
A mature organism is diploid and produces
gametes by meiosis.
Gametes fuse to form diploid zygote; zygote
divides by mitosis to form mature organism.

Figure 7.3 All Sexual Life Cycles Involve Fertilization and Meiosis (Part 3)

Concept 7.1 Different Life Cycles Use Different Modes of Cell

Reproduction

The essence of sexual reproduction is that it

allows the random selection of half the diploid
chromosome set.
This forms a haploid gamete that fuses with
another to make a diploid cell.
Thus, no two individuals have exactly the same
genetic makeup.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Four events must occur for cell division:

Reproductive signalto initiate cell division

Replication of DNA

Segregationdistribution of the DNA into the

two new cells
Cytokinesisdivision of the cytoplasm and
separation of the two new cells

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

In prokaryotes, cell division results in

reproduction of the entire organism.
The cell:
Grows in size
Replicates its DNA

Separates the DNA and cytoplasm into two

cells through binary fission

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Most prokaryotes have one chromosome, a

single molecule of DNAusually circular.
Two important regions in reproduction:
ori - where replication starts
ter - where replication ends

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Replication occurs as the DNA is threaded

through a replication complex of proteins in
the center of the cell.
Replication begins at the ori site and moves
towards the ter site.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

As replication proceeds, the ori complexes move

to opposite ends of the cell.
DNA sequences adjacent to the ori region
actively bind proteins for the segregation,
hydrolyzing ATP for energy.
An actin-like protein provides a filament along
which ori and other proteins move.

Figure 7.4 Prokaryotic Cell Division

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Cytokinesis begins after chromosome

segregation by a pinching in of the plasma
membraneprotein fibers form a ring.
As the membrane pinches in, new cell wall
materials are synthesized resulting in
separation of the two cells.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Eukaryotic cells divide by mitosis followed by

cytokinesis.
Replication of DNA occurs as long strands are
threaded through replication complexes.
DNA replication only occurs during a specific
stage of the cell cycle.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

In segregation of DNA after cell division, one

copy of each chromosome ends up in each of
the two new cells.
In eukaryotes, the chromosomes become highly
condensed.
Mitosis segregates them into two new nuclei
the cytoskeleton is involved in the process.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Cytokinesis follows mitosis.

The process in plant cells (which have cell walls)

is different than in animal cells (which do not
have cell walls).

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

The cell cyclethe period between cell

divisions
In eukaryotes it is divided into mitosis and
cytokinesiscalled the M phaseand a long
interphase.
During interphase, the cell nucleus is visible
and cell functions including replication occur
Interphase begins after cytokinesis and ends
when mitosis starts.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Interphase has three subphases: G1, S, and G2.

G1 (Gap 1)variable, a cell may spend a long

time in this phase carrying out its functions
S phase (Synthesis)DNA is replicated
G2 (Gap 2)the cell prepares for mitosis,
synthesizes microtubules for segregating
chromosomes

Figure 7.5 The Phases of the Eukaryotic Cell Cycle (Part 1)

Figure 7.5 The Phases of the Eukaryotic Cell Cycle (Part 2)

Figure 7.5 The Phases of the Eukaryotic Cell Cycle (Part 3)

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

In mitosis, one nucleus produces two daughter

nuclei each containing the same number of
chromosomes as the parent nucleus.
Mitosis is continuous, but can be can be divided
into phasesprophase, prometaphase,
metaphase, anaphase, and telophase.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

During interphase, only the nuclear envelope

and and the nucleolus are visible.
The chromatin (DNA) is not yet condensed.
Three structures appear in prophase:
The condensed chromosomes

Centrosome
Spindle

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Condensed chromosomes appear during

prophase.
Sister chromatidstwo DNA molecules on
each chromosome after replication

Centromereregion where chromatids are

joined
Kinetochores are protein structures on the
centromeres, and are important for
chromosome movement.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

The karyotype of an organism reflects the

number and sizes of its condensed
chromosomes.
Karyotype analysis can be used to identify
organisms, but DNA sequence is more
commonly used.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Segregation is aided by other structures:

The centrosome determines the orientation of

the spindle apparatus.
Each centrosome can consist of two
centrioleshollow tubes formed by
microtubules.
Centrosome is duplicated during S phase and
each moves towards opposite sides of the
nucleus.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Centrosomes serve as mitotic centers or poles;

the spindle forms between the poles from two
types of microtubules:
Polar microtubules form a spindle and overlap
in the center.
Kinetochore microtubulesattach to
kinetochores on the chromatids. Sister
chromatids attach to opposite halves of the
spindle.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Chromosome separation and movement is

highly organized.
During prometaphase, the nuclear envelope
breaks down.
Chromosomes consisting of two chromatids
attach to the kinetochore mictotubules.

Figure 7.6 The Phases of Mitosis (1)

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Durin metaphase, chromosomes line up at the

midline of the cell.
During anaphase, the separation of sister
chromatids is controlled by M phase cyclinCdk; cohesin is hydrolyzed by separase.
After separation, they move to opposite ends of
the spindle and are referred to as daughter
chromosomes.

Figure 7.6 The Phases of Mitosis (2)

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

A protein at the kinetochorescytoplasmic

dyneinhydrolyzes ATP for energy to move
chromosomes along the microtubules towards
the poles.

Microtubules also shorten, drawing

chromosomes toward poles.

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Telophase occurs after chromosomes have

separated:
Spindle breaks down
Chromosomes uncoil
Nuclear envelope and nucleoli appear
Two daughter nuclei are formed with identical
genetic information

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Cytokinesis:

Division of the cytoplasm differs in plant and

animals
In animal cells, plasma membrane pinches
between the nuclei because of a contractile
ring of microfilaments of actin and myosin.

Figure 7.7 Cytokinesis Differs in Animal and Plant Cells (Part 1)

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

Plant cells:

Vesicles from the Golgi apparatus appear along

the plane of cell division
These fuse to form a new plasma membrane.
Contents of vesicles form the cell platethe
beginning of the new cell wall.

Figure 7.7 Cytokinesis Differs in Animal and Plant Cells (Part 2)

Concept 7.2 Both Binary Fission and Mitosis Produce Genetically

Identical Cells

After cytokinesis:

Each daughter cell contains all of the

components of a complete cell.
Chromosomes are precisely distributed.
The orientation of cell division is important to
development, but organelles are not always
evenly distributed.

Concept 7.3 Cell Reproduction Is Under Precise Control

The reproductive rates of most prokaryotes

respond to environmental conditions.
In eukaryotes, cell division is related to the
needs of the entire organism.
Cells divide in response to extracellular signals,
like growth factors.

Concept 7.3 Cell Reproduction Is Under Precise Control

The eukaryotic cell cycle has four stages: G1, S,

G2, and M.
Progression is tightly regulatedthe G1-S
transition is called R, the restriction point.
Passing this point usually means the cell will
proceed with the cell cycle and divide.

Figure 7.8 The Eukaryotic Cell Cycle

Concept 7.3 Cell Reproduction Is Under Precise Control

Specific signals trigger the transition from one

phase to another.
Evidence for substances as triggers came from
cell fusion experiments.
Nuclei in cells at different stages, fused by
polyethylene glycol, both entered the phase of
DNA replication (S).

Figure 7.9 Regulation of the Cell Cycle (Part 1)

Concept 7.3 Cell Reproduction Is Under Precise Control

Transitions also depend on activation of cyclindependent kinases (Cdks).

A protein kinase is an enzyme that catalyzes
phosphorylation from ATP to a protein.
Phosphorylation changes the shape and function
of a protein by changing its charges.

Concept 7.3 Cell Reproduction Is Under Precise Control

Cdk is activated by binding to cyclin (by

allosteric regulation); this alters its shape and
exposes its active site.
The G1-S cyclin-Cdk complex acts as a protein
kinase and triggers transition from G1 to S.
Other cyclin-Cdks act at different stages of the
cell cycle, called cell cycle checkpoints.

Figure 7.10 Cyclins Are Transient in the Cell Cycle

Concept 7.3 Cell Reproduction Is Under Precise Control

Example of G1-S cyclin-Cdk regulation:

Progress past the restriction point in G1 depends

on retinoblastoma protein (RB).
RB normally inhibits the cell cycle, but when
phosphorylated by G1-S cyclin-Cdk, RB
becomes inactive and no longer blocks the cell
cycle.

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Meiosis consists of two nuclear divisions but

DNA is replicated only once. The function of
meiosis is to:
Reduce the chromosome number from diploid
to haploid
Ensure that each haploid has a complete set of
chromosomes

Generate diversity among the products

Figure 7.11 Mitosis and Meiosis: A Comparison

Figure 7.12 Meiosis: Generating Haploid Cells (Part 1)

Figure 7.12 Meiosis: Generating Haploid Cells (Part 2)

Figure 7.12 Meiosis: Generating Haploid Cells (Part 3)

Figure 7.12 Meiosis: Generating Haploid Cells (Part 4)

Figure 7.12 Meiosis: Generating Haploid Cells (Part 5)

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Meiotic division reduces the chromosome

number. Two unique features:
In meiosis I, homologous pairs of
chromosomes come together and line up along
their entire lengths.
After metaphase I, the homologous
chromosome pairs separate, but individual
chromosomes made up of two sister
chromatids remain together.

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Meiosis I is preceded by an S phase during

which DNA is replicated.
Each chromosome then consists of two sister
chromatids, held together by cohesin proteins.
At the end of meiosis I, two nuclei form, each
with half the original chromosomesstill
composed of sister chromatids.

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Sister chromatids separate during meiosis II,

which is not proceeded by DNA replication.
The products of meiosis I and II are four cells
with a haploid number of chromosomes.
These four cells are not genetically identical.
Two processes may occur:
Crossing over and independent assortment

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

In prophase of meiosis I homologous

chromosomes pair by synapsis.
The four chromatids of each pair of
chromosomes form a tetrad,or bivalent.
The homologs seem to repel each other but are
held together at chiasmata.

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Crossing over is an exchange of genetic

material that occurs at the chiasma.
Crossing over results in recombinant
chromatids and increases genetic variability of
the products.

In-Text Art, Ch. 7, p. 138

Figure 7.13 Crossing Over Forms Genetically Diverse Chromosomes

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Prophase I may last a long time.

Human males: Prophase I lasts about 1 week,

and 1 month for entire meiotic cycle
Human females: Prophase I begins before
birth, and ends up to decades later during the
monthly ovarian cycle

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Independent assortment during anaphase I

also allows for chance combinations and
genetic diversity.
After homologous pairs of chromosomes line up
at metaphase I, it is a matter of chance which
member of a pair goes to which daughter cell.
The more chromosomes involved, the more
combinations possible.

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Meiotic errors:

Nondisjunctionhomologous pairs fail to

separate at anaphase Isister chromatids fail
to separate, or homologous chromosomes may
not remain together
Either results in aneuploidychromosomes
lacking or present in excess

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

Organisms with triploid (3n), tetraploid (4n), and

even higher levels are called polyploid.
This can occur through an extra round of DNA
duplication before meiosis, or the lack of
spindle formation in meiosis II.
Polyploidy occurs naturally in some species,
and can be desirable in plants.

Concept 7.4 Meiosis Halves the Nuclear Chromosome Content

and Generates Diversity

If crossing over happens between nonhomologous chromosomes, the result is a

translocation.
A piece of chromosome may rejoin another
chromosome, and its location can have
profound effects on the expression of other
genes.

Example: Leukemia

In-Text Art, Ch. 7, p. 140

Concept 7.5 Programmed Cell Death Is a Necessary Process in

Living Organisms

Cell death occurs in two ways:

In necrosis, the cell is damaged or starved for

oxygen or nutrients. The cell swells and bursts.
Cell contents are released to the extracellular
environment and can cause inflammation.

7.5 ProConcept 7.ammed Cell Death Is a Necessary Process in

Living Organisms

Apoptosis is genetically programmed cell

death. Two possible reasons:
The cell is no longer needed, e.g., the
connective tissue between the fingers of a
fetus.
Old cells may be prone to genetic damage that
can lead to cancerblood cells and epithelial
cells die after days or weeks.

Concept 7.5 Programmed Cell Death Is a Necessary Process in

Living Organisms

Events of apoptosis:

Cell detaches from its neighbors

Cuts up its chromatin into nucleosome-sized
pieces
Forms membranous lobes called blebs that
break into fragments
Surrounding living cells ingest the remains of
the dead cell

Figure 7.14 Apoptosis: Programmed Cell Death (Part 1)

Concept 7.5 Programmed Cell Death Is a Necessary Process in

Living Organisms

Cell death cycle is controlled by signals:

Lack of a mitotic signal (growth factor)

Recognition of damaged DNA

External signals cause membrane proteins to

change shape and activate enzymes called
caspaseshydrolyze proteins of membranes.

Figure 7.14 Apoptosis: Programmed Cell Death (Part 2)

Answer to Opening Question

Human papilloma virus (HPV) stimulates the

cell cycle when it infects the cervix.

Two proteins regulate the cell cycle:

Oncogene proteins are positive regulators
of the cell cyclein cancer cells they are
overactive or present in excess
Tumor suppressors are negative
regulators of the cell cycle, but in cancer
cells they are inactivecan be blocked by
a virus such as HPV

Figure 7.15 Molecular Changes Regulate the Cell Cycle in Cancer Cells