Chapter 11

Cell
Communication

Lecture Presentations by
Nicole Tunbridge and
© 2017 Pearson Education, Inc.
Kathleen Fitzpatrick
Cellular Messaging

▪Cells can signal to

each other and
interpret the signals
they receive from other
cells and the
environment
Cellular Messaging

▪Signals are most often

chemicals
▪The same small set of cell-
signaling mechanisms
shows up in diverse species
and processes
Figure 11.1

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Concept 11.1: External signals are converted to
responses within the cell

▪Communication among
microorganisms provides
some insight into how
cells send, receive, and
respond to signals
Evolution of Cell Signaling

▪The yeast Saccharomyces

cerevisiae has two mating
types, a and α
▪Cells of different mating
types locate each other via
secreted factors specific to
each type
Evolution of Cell Signaling

▪ The binding of a mating factor

at the cell surface initiates a
series of steps called a signal
transduction pathway
▪ Molecular details of signal
transduction in yeasts and
mammals are very similar.
Figure 11.2_1

Receptor α factor
1 Exchange of
mating factors

a α

Yeast cell, a factor Yeast cell,

mating type a mating type α

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Figure 11.2_2

Receptor α factor
1 Exchange of
mating factors

a α

Yeast cell, a factor Yeast cell,

mating type a mating type α

2 Mating

a α

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Figure 11.2_3

Receptor α factor
1 Exchange of
mating factors

a α

Yeast cell, a factor Yeast cell,

mating type a mating type α

2 Mating

a α

3 New a/α cell

a/α

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▪Ancestral signaling
molecules likely evolved
in prokaryotes and
single-celled eukaryotes
and were adopted for use
in their multicellular
descendants
▪Cell signaling is critical
among prokaryotes
▪A concentration of signaling
molecules allows bacteria to
sense local population
density in a process called
quorum sensing
Figure 11.3

1 Individual
rod-shaped
cells

2 Aggregation
in progress
0.5 mm

3 Spore-forming
structure
2.5 mm
(fruiting body)

Fruiting bodies
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▪An example of quorum
sensing is the formation
of a biofilm
▪A biofilm is an
aggregation of bacterial
cells adhered to a
surface
▪ Another example of medical
importance is the secretion of
toxins by infectious bacteria
▪ Interfering with the signaling
pathways used in quorum
sensing may be a promising
approach as an alternative to
antibiotic treatment
Local and Long-Distance Signaling

▪Cells in a multicellular
organism communicate
via signaling molecules
▪In local signaling, animal
cells may communicate
by direct contact
Local and Long-Distance Signaling

▪Animal and plant cells have

cell junctions that directly
connect the cytoplasm of
adjacent cells
▪Signaling substances in the
cytosol can pass freely
between adjacent cells
Figure 11.4
Plasma membranes Cell wall

Gap junctions Plasmodesmata

between animal cells between plant cells
(a) Cell junctions

(b) Cell-cell recognition

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▪In other cases, animal
cells communicate
using secreted
messenger molecules
that travel only short
distances
▪Growth factors, which
stimulate nearby target cells
to grow and divide, are one
class of such local
regulators in animals
▪This type of local signaling
in animals is called
paracrine signaling
▪Synaptic signaling
occurs in the animal
nervous system when
a neurotransmitter is
released in response
to an electric signal
▪Local signaling in
plants is not well
understood beyond
communication
between
plasmodesmata
▪In long-distance
signaling, plants
and animals use
chemicals called
hormones
▪Hormonal signaling in
animals is called
endocrine signaling;
specialized cells release
hormones, which travel
to target cells via the
circulatory system
▪The ability of a cell to
respond to a signal
depends on whether or
not it has a receptor
specific to that signal
Figure 11.5
Local signaling
Target cells Electrical signal triggers
release of neurotransmitter.
Neurotransmitter
diffuses across
Signaling synapse.
cell

Secretory
vesicles
Local regulator Target cell
(a) Paracrine signaling (b) Synaptic signaling

Long-distance signaling
Endocrine cell Target cell
specifically
binds
hormone.

Hormone
travels in
bloodstream.
Blood
vessel
(c) Endocrine (hormonal) signaling
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The Three Stages of Cell Signaling: A Preview

▪Earl W. Sutherland
and colleagues
discovered how the
hormone epinephrine
acts on cells
The Three Stages of Cell Signaling: A Preview

▪Sutherland suggested that

cells receiving signals went
through three processes
▪Reception
▪Transduction
▪Response
▪In reception, the
target cell detects a
signaling molecule
that binds to a
receptor protein on
the cell surface
▪In transduction, the binding
of the signaling molecule
alters the receptor and
initiates a signal
transduction pathway;
transduction often occurs in
a series of steps
▪In response, the
transduced signal
triggers a specific
response in the
target cell
Figure 11.6_1

EXTRACELLULAR CYTOPLASM
FLUID Plasma membrane
1 Reception

Receptor

Signaling
molecule

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Figure 11.6_2

EXTRACELLULAR CYTOPLASM
FLUID Plasma membrane
1 Reception 2 Transduction

Receptor

1 2 3
Relay molecules

Signaling
molecule

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Figure 11.6_3

EXTRACELLULAR CYTOPLASM
FLUID Plasma membrane
1 Reception 2 Transduction 3 Response

Receptor
Activation
1 2 3 of cellular
response
Relay molecules

Signaling
molecule

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Concept 11.2: Reception: A signaling molecule
binds to a receptor protein, causing it to change
shape

▪ The binding between a signal

molecule (ligand) and receptor
is highly specific
▪ A shape change in a receptor is
often the initial transduction of
the signal
Concept 11.2: Reception: A signaling molecule
binds to a receptor protein, causing it to change
shape

▪Most signal
receptors are
plasma membrane
proteins
Receptors in the Plasma Membrane

▪G protein-coupled
receptors (GPCRs)
are the largest family
of cell-surface
receptors
Receptors in the Plasma Membrane

▪Most water-soluble
signal molecules bind
to specific sites on
receptor proteins that
span the plasma
membrane
Figure 11.7

β2-adrenergic Molecule
receptors that mimics
ligand

Plasma membrane Cholesterol

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▪There are three main
types of membrane
receptors:
▪G protein-coupled
receptors
▪Receptor tyrosine kinases
▪Ion channel receptors
▪G protein-coupled
receptors (GPCRs) are
cell-surface transmembrane
receptors that work with the
help of a G protein
▪G proteins bind the energy-
rich GTP
▪G proteins are all very
similar in structure
▪GPCR systems are
extremely widespread
and diverse in their
functions
Figure 11.8a

Signaling molecule binding site

Segment that
interacts with
G proteins inside the cell
G protein-coupled receptor

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▪Receptor tyrosine
kinases (RTKs) are
membrane receptors
that transfer phosphate
groups from ATP to
another protein
▪A receptor tyrosine
kinase can trigger
multiple signal
transduction
pathways at once
▪Abnormal functioning
of RTKs is associated
with many types of
cancers
Figure 11.8b

G protein-coupled Plasma membrane Activated Signaling

receptor receptor molecule

GTP
GDP
G protein Inactive
GDP GTP
(inactive) Enzyme enzyme
1 2
Activated EXTRACELLULAR
enzyme

GTP GDP

P
i
Cellular
response CYTOPLASM
3 4

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▪A ligand-gated ion
channel receptor acts
as a gate that opens
and closes when the
receptor changes
shape
▪When a signal molecule
binds as a ligand to the
receptor, the gate allows
specific ions, such as
+ 2+
Na or Ca , through a
channel in the receptor
Figure 11.8c
Signaling molecule Signaling molecule EXTRACELLULAR
(ligand) Ligand-binding site
α helix in the
membrane

Tyr Tyr Tyr Tyr Tyr Tyr

Tyrosines Tyr Tyr Tyr Tyr Tyr Tyr
Tyr Tyr Tyr Tyr Tyr Tyr

Receptor tyrosine Dimer

kinase proteins
(inactive monomers) CYTOPLASM
1 2

Activated
relay proteins

Cellular
P Tyr Tyr P
Tyr Tyr P Tyr Tyr P response 1
P Tyr Tyr P
Tyr Tyr P Tyr Tyr P
P Tyr Tyr P
Tyr Tyr
6 ATP 6 ADP
P Tyr Tyr P Cellular
response 2
Activated tyrosine Fully activated receptor
kinase regions tyrosine kinase
(unphosphorylated (phosphorylated Inactive
dimer) dimer) relay proteins

3 4
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Intracellular Receptors
▪ Intracellular receptor proteins
are found in the cytoplasm or
nucleus of target cells
▪ Small or hydrophobic chemical
messengers can readily cross
the membrane and activate
receptors
Intracellular Receptors
▪ Examples of hydrophobic
messengers are the steroid and
thyroid hormones of animals
▪ An activated hormone-receptor
complex can act as a
transcription factor, turning on
or off specific genes
Figure 11.9
Hormone EXTRA-
(aldosterone) CELLULAR
FLUID

Plasma
Receptor membrane
protein
Hormone-
receptor
complex

DNA

mRNA
New
NUCLEUS protein

CYTOPLASM
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Concept 11.3: Transduction: Cascades of
molecular interactions relay signals from
receptors to target molecules in the cell

▪Cell signaling is
usually a multistep
process
▪Multistep pathways
can greatly amplify a
Concept 11.3: Transduction: Cascades of
molecular interactions relay signals from
receptors to target molecules in the cell

▪Multistep pathways provide

more opportunities for
coordination and regulation
of the cellular response
Signal Transduction Pathways

▪The binding of a
signaling molecule to a
receptor triggers the
first step in a chain of
molecular interactions
Signal Transduction Pathways

▪The receptor activates

another protein, which
activates another, and so
on, until the protein
producing the response
is activated
Signal Transduction Pathways

▪At each step, the

signal is transduced
into a different form,
usually a shape
change in a protein
Protein Phosphorylation and
Dephosphorylation

▪Phosphorylation and
dephosphorylation of
proteins is a widespread
cellular mechanism for
regulating protein activity
Protein Phosphorylation and
Dephosphorylation

▪Protein kinases
transfer phosphates
from ATP to protein, a
process called
phosphorylation
Protein Phosphorylation and
Dephosphorylation

▪Many relay molecules in

signal transduction
pathways are protein
kinases, creating a
phosphorylation
cascade
Figure 11.10

Signaling molecule

Activated relay
Receptor molecule

Inactive
protein kinase
1
Active
protein
kinase
1
Inactive
protein kinase
2 ATP
ADP
Active P
protein
PP kinase
Pi 2

Inactive
protein ATP
ADP P
Active
Cellular
protein
PP response
Pi

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▪Protein phosphatases
rapidly remove the
phosphates from
proteins, a process called
dephosphorylation
▪This phosphorylation and
dephosphorylation
system acts as a
molecular switch, turning
activities on and off or up
or down, as required
Small Molecules and Ions as Second
Messengers
▪ Many signaling pathways
involve second messengers
▪ These are small, nonprotein,
water-soluble molecules or ions
that spread throughout a cell by
diffusion
Small Molecules and Ions as Second
Messengers
▪Second messengers
participate in pathways
initiated by GPCRs and
RTKs
▪Cyclic AMP and calcium
ions are common second
messengers
Cyclic AMP

▪ Cyclic AMP (cAMP) is one of

the most widely used second
messengers
▪ Adenylyl cyclase, an enzyme
in the plasma membrane,
converts ATP to cAMP in
response to an extracellular
signal
Figure 11.11

5′C
P P P Adenylyl cyclase Phosphodiesterase P
P
Pyrophosphate 3′C H2O
ATP P Pi
cAMP AMP

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▪Many signal molecules
trigger formation of cAMP
▪Other components of
cAMP pathways are G
proteins, G protein-
coupled receptors, and
protein kinases
▪cAMP usually activates
protein kinase A, which
phosphorylates various
other proteins
▪Further regulation of cell
metabolism is provided by G
protein systems that inhibit
adenylyl cyclase
Figure 11.12

First messenger
G protein- (signaling molecule
coupled such as epinephrine) Adenylyl
receptor 1 G protein cyclase
(GPCR)

2 GTP 3

ATP 4 Second
messenger
cAMP

Protein
5 kinase A

Cellular responses

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▪ Understanding of the role of
cAMP in G protein signaling
pathways helps explain how
certain microbes cause disease
▪ The cholera bacterium, Vibrio
cholerae, produces a toxin that
modifies a G protein so that it is
stuck in its active form
▪ This protein continually makes
cAMP, causing intestinal cells
to secrete large amounts of salt
into the intestines
▪ Water follows by osmosis, and
an untreated person can soon
die from loss of water and salt
Calcium Ions and Inositol Triphosphate (IP3)

▪Calcium ions 2+
(Ca )
are used widely as a
second messenger
Calcium Ions and Inositol Triphosphate (IP3)

▪Ca2+ can function as a

second messenger because
its concentration in the
cytosol is normally much
lower than the concentration
outside the cell
Calcium Ions and Inositol Triphosphate (IP3)

▪A small change in
number of calcium ions
thus represents a
relatively large
percentage change in
calcium concentration
Figure 11.13

Endoplasmic Plasma
reticulum (ER) membrane
ATP

Mitochondrion

Nucleus

Ca2+
pump

CYTOSOL ATP

EXTRACELLULAR
FLUID

High [Ca2+] Low [Ca2+]

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▪A signal relayed by a
signal transduction
pathway may trigger
an increase in
calcium in the
cytosol
▪Pathways leading to the
release of calcium
involve inositol
triphosphate (IP3) and
diacylglycerol (DAG) as
additional second
messengers
▪These two are
produced by
cleavage of a certain
phospholipid in the
plasma membrane
Figure 11.14_1

EXTRA- Signaling molecule

CELLULAR (first messenger)
FLUID
G protein

GTP
CYTOSOL G protein-coupled DAG
receptor Phospholipase C
PIP2
IP3-gated
calcium channel IP3
Endoplasmic
(second messenger)
reticulum (ER)
lumen
Ca2+

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Concept 11.4: Response: Cell signaling leads to
regulation of transcription or cytoplasmic
activities

▪The cell’s response

to an extracellular
signal is called the
“output response”
Nuclear and Cytoplasmic Responses

▪Ultimately, a signal
transduction pathway leads
to regulation of one or more
cellular activities
▪The response may occur in
the nucleus or in the
cytoplasm
Nuclear and Cytoplasmic Responses

▪ Many signaling pathways

regulate the synthesis of
enzymes or other proteins,
usually by turning genes on or
off in the nucleus
▪ The final activated molecule in
the signaling pathway may
function as a transcription factor
Figure 11.15
Growth factor Reception
Receptor

Phospho-
rylation Transduction
cascade
CYTOPLASM

Inactive Active
transcription transcription
factor factor Response
P
DNA

Gene

NUCLEUS mRNA
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▪Other pathways may
regulate the activity
of enzymes rather
than their synthesis
▪For example, a signal
could cause opening or
closing of an ion channel
in the plasma membrane
or a change in cell
metabolism
Figure 11.16

Reception Transduction
Binding of epinephrine to G protein-coupled Inactive
receptor G protein
(1 molecule)
Active G protein (102 molecules)
Inactive
adenylyl cyclase
Active adenylyl cyclase (102)

ATP
Cyclic AMP (104)
Inactive
protein kinase A
Active protein kinase A (104)
Inactive
phosphorylase kinase
Response Active phosphorylase kinase (105)
Inactive
Glycogen
glycogen phosphorylase
Glucose 1-phosphate
(108 molecules) Active glycogen phosphorylase (106)

© 2017 Pearson Education, Inc.

▪Signaling pathways
can also affect the
overall behavior of a
cell; for example, a
signal could lead to
cell division
Regulation of the Response

▪A response to a signal
may not be simply “on”
or “off”
▪There are four aspects
of signal regulation:
Regulation of the Response
▪Amplification of the signal
(and thus the response)
▪Specificity of the response
▪Overall efficiency of response,
enhanced by scaffolding
proteins
▪Termination of the signal
Signal Amplification

▪Enzyme cascades amplify

the cell’s response to the
signal
▪At each step, the number
of activated products can
be much greater than in
the preceding step
The Specificity of Cell Signaling and
Coordination of the Response
▪Different kinds of cells have
different collections of
proteins
▪These different proteins
allow cells to detect and
respond to different signals
The Specificity of Cell Signaling and
Coordination of the Response
▪ The same signal can have
different effects in cells with
different proteins and pathways
▪ Pathway branching and “cross-
talk” further help the cell
coordinate incoming signals
Figure 11.17

Signaling
molecule

Receptor

Relay
mole-
Activation
cules
or inhibition

Response 1 Response 2 Response 3 Response 4 Response 5

Cell A: Pathway leads Cell B: Pathway Cell C: Cross-talk Cell D: Different

to a single response. branches, leading to occurs between two receptor leads to a
two responses. pathways. different response.

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Signaling Efficiency: Scaffolding Proteins and
Signaling Complexes

▪Scaffolding proteins
are large relay
proteins to which
other relay proteins
are attached
Signaling Efficiency: Scaffolding Proteins and
Signaling Complexes

▪Scaffolding proteins can

increase the signal
transduction efficiency by
grouping together different
proteins involved in the
same pathway
Signaling Efficiency: Scaffolding Proteins and
Signaling Complexes

▪In some cases,

scaffolding proteins
may also help activate
some of the relay
proteins
Figure 11.18

Signaling Plasma
molecule membrane

Receptor

Three
different
protein
kinases
Scaffolding
protein

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Termination of the Signal

▪Inactivation
mechanisms are an
essential aspect of
cell signaling
Termination of the Signal

▪If the concentration

of external signaling
molecules falls,
fewer receptors will
be bound
Termination of the Signal

▪Unbound
receptors revert to
an inactive state
Concept 11.5: Apoptosis integrates multiple
cell-signaling pathways

▪ Cells that are infected,

damaged, or at the end of their
functional lives often undergo
“programmed cell death”
▪ Apoptosis is the best-
understood type
Concept 11.5: Apoptosis integrates multiple
cell-signaling pathways

▪ Components of the cell are

chopped up and packaged into
vesicles that are digested by
scavenger cells
▪ Apoptosis prevents enzymes
from leaking out of a dying cell
and damaging neighboring cells
Figure 11.19

2 µm

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Apoptosis in the Soil Worm Caenorhabditis
elegans
▪In worms and other
organisms, apoptosis is
triggered by signals that
activate a cascade of
“suicide” proteins in the cells
programmed to die
Apoptosis in the Soil Worm Caenorhabditis
elegans
▪When the death signal is
received, an apoptosis-
inhibiting protein (Ced-9) is
inactivated, triggering a
cascade of caspase
proteins that promote
apoptosis
Apoptosis in the Soil Worm Caenorhabditis
elegans

▪The chief caspase in

the nematode is called
Ced-3
Figure 11.20

Active Ced-9 protein Inactive Ced-9 does Cell

inhibits Ced-4 activity. not inhibit Ced-4. forms
blebs
Mitochondrion Death-
signaling
molecule

Active Active
Ced-4 Ced-3 Nucleases

Other
Ced-4 Ced-3 proteases
Receptor Activation
for death- Inactive proteins cascade
signaling
molecule
(a) No death signal (b) Death signal

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Apoptotic Pathways and the Signals That
Trigger Them

▪In humans and other

mammals, several
different pathways,
including about 15
caspases, can carry out
apoptosis
Apoptotic Pathways and the Signals That
Trigger Them

▪Apoptosis can be triggered

by signals from outside the
cell or inside it
▪Internal signals can result
from irreparable DNA
damage or excessive
protein misfolding
▪Apoptosis evolved early
in animal evolution and is
essential for the
development and
maintenance of all
animals
▪For example,
apoptosis is a normal
part of development of
hands and feet in
humans (and paws in
other mammals)
▪Apoptosis may be involved
in some diseases (for
example, Parkinson’s and
Alzheimer’s); interference
with apoptosis may
contribute to some cancers
Figure 11.21

Interdigital Cells undergoing Space between

tissue apoptosis 1 mm digits

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 Chapter 12

The Cell Cycle

Lecture Presentations by
Nicole Tunbridge and
© 2017 Pearson Education, Inc.
Kathleen Fitzpatrick
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 12.1

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▪In unicellular organisms,
division of one cell
reproduces the entire
organism
▪Multicellular eukaryotes
depend on cell division
for
 ▪development from a fertilized
egg
▪growth
▪repair
▪Cell division is an integral
part of the cell cycle, the
life of a cell from formation
to its own division
Figure 12.2
100 µm (a) Asexual reproduction

50 µm

(b) Growth and

development

(c) Tissue renewal

20 µm
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Concept 12.1: Most cell division results in
genetically identical daughter cells

▪Most cell division

results in two daughter
cells with identical
genetic information,
DNA
Concept 12.1: Most cell division results in
genetically identical daughter cells

▪The exception is
meiosis, a special type
of division that can
produce sperm and
egg cells
Cellular Organization of the Genetic Material

▪All the DNA in a

cell constitutes
the cell’s
genome
Cellular Organization of the Genetic Material

▪A genome can consist of

a single DNA molecule
(common in prokaryotic
cells) or a number of
DNA molecules (common
in eukaryotic cells)
Cellular Organization of the Genetic Material

▪DNA molecules
in a cell are
packaged into
chromosomes
Figure 12.3

20 µm

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▪ 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
Distribution of Chromosomes During
Eukaryotic Cell Division

▪In preparation for cell

division, DNA is
replicated and the
chromosomes
condense
Distribution of Chromosomes During
Eukaryotic Cell Division

▪Each duplicated
chromosome has two sister
chromatids (joined copies
of the original
chromosome), attached
along their lengths by
cohesins
Distribution of Chromosomes During
Eukaryotic Cell Division

▪The centromere is the

narrow “waist” of the
duplicated chromosome,
where the two
chromatids are most
closely attached
Figure 12.4

Sister
chromatids

Centromeres, one on
each sister chromatid 0.5 µm

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▪During cell division, the two
sister chromatids of each
duplicated chromosome
separate and move into two
nuclei
▪Once separate, the
chromatids are called
chromosomes
Figure 12.5_1
1 Chromosomes Chromosomal
DNA molecules
Centromere

Chromosome
arm

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Figure 12.5_2
1 Chromosomes Chromosomal
DNA molecules
Centromere

Chromosome
arm
Chromosome duplication

Sister
chromatids

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Figure 12.5_3
1 Chromosomes Chromosomal
DNA molecules
Centromere

Chromosome
arm
Chromosome duplication

Sister
chromatids
Separation of sister
chromatids
3

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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 are produced by a
variation of cell division
called meiosis
▪Meiosis yields nonidentical
daughter cells that have half
as many chromosomes as
the parent 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
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)
▪Interphase (about 90% of
the cell cycle) can be
divided into three
phases:
▪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
Figure 12.6

S
G1
(DNA synthesis)

G2

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▪Mitosis is conventionally
broken down into five
stages:
▪prophase
▪prometaphase
▪metaphase
▪anaphase
▪telophase
Figure 12.7a

10 µm
G2 of Interphase Prophase Prometaphase
Centrosomes Chromosomes Early mitotic Fragments Nonkinetochore
(with centriole (duplicated, Aster of nuclear
spindle microtubules
pairs) uncondensed) Centromere envelope

Plasma
Nucleolus Two sister chromatids Kinetochore Kinetochore
Nuclear membrane
of one chromosome microtubules
envelope
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Figure 12.7b

10 µm
Metaphase Anaphase Telophase and Cytokinesis
Metaphase Cleavage Nucleolus
plate furrow forming

Daughter
chromosomes
Spindle Nuclear
Centrosome at envelope
one spindle pole forming
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The Mitotic Spindle: A Closer Look

▪The mitotic spindle is

a structure made of
microtubules that
controls chromosome
movement during
mitosis
The Mitotic Spindle: A Closer Look

▪In animal cells,

assembly of spindle
microtubules begins in
the centrosome, the
microtubule-organizing
center
The Mitotic Spindle: A Closer Look

▪The centrosome
replicates during
interphase, forming two
centrosomes that migrate
to opposite ends of the
cell during prophase and
prometaphase
▪An aster (a radial
array of short
microtubules)
extends from each
centrosome
▪The spindle includes
the centrosomes, the
spindle microtubules,
and the asters
▪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, a
plane midway between
the spindle’s two poles
Figure 12.8
Aster Centrosome
Sister
chromatids Metaphase
plate
(imaginary)

Kineto-
chores

Microtubules
Overlapping
nonkinetochore
microtubules
Kinetochore
microtubules
Chromosomes

Centrosome

1 µm 0.5 µm
© 2017 Pearson Education, Inc.
▪In anaphase the
cohesins are
cleaved by an
enzyme called
separase
▪Sister chromatids
separate and move
along the kinetochore
microtubules toward
opposite ends of the
cell
▪The microtubules
shorten by
depolymerizing at
their kinetochore
ends
▪Results of a clever
experiment suggest that
motor proteins on
kinetochores “walk” the
chromosomes along the
microtubules during
anaphase
▪The depolymerization of
the microtubules at the
kinetochore ends occurs
after the motor proteins
have passed
▪This is called the “Pac-
man” mechanism
▪Nonkinetochore
microtubules from
opposite poles overlap
and push against each
other, elongating the
cell
▪At the end of
anaphase, duplicate
groups of
chromosomes have
arrived at opposite
ends of the elongated
cell
▪Cytokinesis begins
during anaphase or
telophase, and the
spindle eventually
disassembles
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
Figure 12.10
(a) Cleavage of an animal cell (SEM) (b) Cell plate formation in a plant cell
(TEM)

Cleavage furrow 100 µm Vesicles Wall of parent cell 1 µm

forming New cell
cell plate Cell plate wall

Contractile ring of Daughter cells

microfilaments Daughter cells
© 2017 Pearson Education, Inc.
Binary Fission in Bacteria

▪Prokaryotes (bacteria
and archaea)
reproduce by a type of
cell division called
binary fission
Binary Fission in Bacteria

▪In binary fission, the

chromosome replicates
(beginning at the origin of
replication), and the two
daughter chromosomes
actively move apart
Binary Fission in Bacteria

▪The plasma
membrane pinches
inward, dividing the
cell into two
Figure 12.12_1
Origin of Cell wall
replication Plasma
membrane
Bacterial cell Bacterial
1 Chromosome
replication Two copies chromosome
begins. of origin

© 2017 Pearson Education, Inc.

Figure 12.12_2
Origin of Cell wall
replication Plasma
membrane
Bacterial cell Bacterial
1 Chromosome
replication Two copies chromosome
begins. of origin

Origin Origin
2 One copy of the
origin is now at
each end of the
cell.

© 2017 Pearson Education, Inc.

Figure 12.12_3
Origin of Cell wall
replication Plasma
membrane
Bacterial cell Bacterial
1 Chromosome
replication Two copies chromosome
begins. of origin

Origin Origin
2 One copy of the
origin is now at
each end of the
cell.

3 Replication
finishes.

© 2017 Pearson Education, Inc.

Figure 12.12_4
Origin of Cell wall
replication Plasma
membrane
Bacterial cell Bacterial
1 Chromosome
replication Two copies chromosome
begins. of origin

Origin Origin
2 One copy of the
origin is now at
each end of the
cell.

3 Replication
finishes.

4 Two daughter
cells result.
© 2017 Pearson Education, Inc.
The Evolution of Mitosis

▪Because prokaryotes
evolved before
eukaryotes, mitosis
probably evolved from
binary fission
The Evolution of Mitosis

▪Certain protists exhibit

types of cell division
that seem intermediate
between binary fission
and mitosis
Figure 12.13

Bacterial Kinetochore
chromosome microtubule

Intact nuclear
envelope

(a) Bacteria (c) Diatoms and some yeasts

Chromosomes
Kinetochore
Microtubules microtubule

Intact nuclear Fragments of

envelope nuclear
envelope
(b) Dinoflagellates (d) Most eukaryotes

© 2017 Pearson Education, Inc.

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
The Cell Cycle Control System

▪The cell cycle

appears to be driven
by specific chemical
signals present in the
cytoplasm
The Cell Cycle Control System
▪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
Figure 12.14
Experiment Experiment 1 Experiment 2

S G1 M G1

Results

S S M M
G1 nucleus G1 nucleus began
immediately entered mitosis without
S phase and DNA chromosome
was synthesized. duplication.

Conclusion Molecules present in the cytoplasm control

the progression to S and M phases.
Data from R. T. Johnson and P. N. Rao, Mammalian cell fusion: Induction of premature
chromosome condensation in interphase nuclei, Nature 226:717–722 (1970).
© 2017 Pearson Education, Inc.
▪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
Figure 12.15
G1 checkpoint

Control
G1 system S

M G2

M checkpoint
G2 checkpoint
© 2017 Pearson Education, Inc.
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 Cell Cycle Clock: Cyclins and Cyclin-
Dependent Kinases

▪The activity of a Cdk

rises and falls with
changes in
concentration of its
cyclin partner
The Cell Cycle Clock: Cyclins and Cyclin-
Dependent Kinases

▪MPF (maturation-
promoting factor) is a
cyclin-Cdk complex that
triggers a cell’s passage
past the G2 checkpoint
into the M phase
Figure 12.16

M G1 S G2 M G1 S G2 M G1
Cdk
MPF Cyclin
activity concentration
Degraded
cyclin Cdk
Cyclin is
degraded
MPF Cyclin
Time G2
checkpoint
(a) Fluctuation of MPF activity and cyclin (b) Molecular mechanisms that help
concentration during the cell cycle regulate the cell cycle

© 2017 Pearson Education, Inc.

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

▪Many signals registered

at checkpoints come
from cellular surveillance
mechanisms within the
cell
Stop and Go Signs: Internal and External
Signals at the Checkpoints

▪Checkpoints also register

signals from outside the
cell
▪Three important
checkpoints are those in
the G1, G2, and M phases
▪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
Figure 12.17
G1 checkpoint

G0

G1 G1
Without go-ahead signal, cell With go-ahead signal, cell
G1 enters G0. continues cell cycle.
S
(a) G1 checkpoint
M G2
G1 G1

M G2 M G2

M checkpoint

G2
Anaphase checkpoint

Prometaphase Metaphase
Without full chromosome attachment, With full chromosome
stop signal is received. attachment, go-ahead signal is
received.
(b) M checkpoint
© 2017 Pearson Education, Inc.
▪An example of an internal
signal is that cells will not
begin anaphase until all
chromosomes are
properly attached to the
spindle at the metaphase
plate
▪This mechanism
ensures that
daughter cells have
the correct number
of chromosomes
▪External factors that
influence cell division
include specific growth
factors
▪Growth factors are
released by certain cells
and stimulate other cells to
divide
▪Platelet-derived growth
factor (PDGF) is made by
blood cell fragments
called platelets
▪In density-dependent
inhibition, crowded cells
will stop dividing
Figure 12.18_1
Scalpels
1 A sample of
human connective
tissue is cut
up into small
pieces. Petri
dish

© 2017 Pearson Education, Inc.

Figure 12.18_2
Scalpels
1 A sample of
human connective
tissue is cut
up into small
pieces. Petri
dish

2 Enzymes digest
the extracellular
matrix, resulting
in a suspension of
free fibroblasts.

© 2017 Pearson Education, Inc.

Figure 12.18_3
Scalpels
1 A sample of
human connective
tissue is cut
up into small
pieces. Petri
dish

2 Enzymes digest
the extracellular
matrix, resulting
in a suspension of
free fibroblasts.

3 Cells are transferred

to culture vessels.

© 2017 Pearson Education, Inc.

Figure 12.18_4
Scalpels
1 A sample of
human connective
tissue is cut
up into small
pieces. Petri
dish

2 Enzymes digest
the extracellular
matrix, resulting
in a suspension of
free fibroblasts.

3 Cells are transferred

to culture vessels. 4 PDGF is added
to half the
vessels.

10 µm
Without PDGF With PDGF Cultured fibroblasts (SEM)
© 2017 Pearson Education, Inc.
▪Most cells also exhibit
anchorage
dependence—to
divide, they must be
attached to a
substratum
▪Density-dependent
inhibition and
anchorage
dependence check the
growth of cells at an
optimal density
▪Cancer cells
exhibit neither
type of regulation
of their division
Figure 12.19

Anchorage dependence: cells

require a surface for division

Density-dependent inhibition:
cells form a single layer

Density-dependent inhibition:
cells divide to fill a gap and
then stop

20 µm 20 µm

(a) Normal mammalian cells (b) Cancer cells

© 2017 Pearson Education, Inc.

Loss of Cell Cycle Controls in Cancer Cells

▪Cancer cells do not respond

normally to the body’s
control mechanisms
▪Cancer cells do not need
growth factors to grow and
divide:
Loss of Cell Cycle Controls in Cancer Cells
▪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
▪Cells that acquire the
ability to divide
indefinitely are
undergoing
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
undergo metastasis, the
spread of cancer cells to
other parts of the body,
where they may form
additional tumors
▪Localized tumors
may be treated with
high-energy
radiation, which
damages the DNA in
the cancer cells
▪To treat metastatic
cancers,
chemotherapies that
target the cell cycle
may be used
Figure 12.20

5 µm
Breast cancer cell
(colorized SEM)

Metastatic
Lymph tumor
vessel
Tumor
Blood
vessel

Glandular Cancer
tissue cell

1 A tumor grows 2 Cancer cells invade 3 Cancer cells spread 4 A small percentage
from a single neighboring tissue. through lymph and of cancer cells may
cancer cell. blood vessels to other metastasize to
parts of the body. another part of the
body.

© 2017 Pearson Education, Inc.

▪Recent advances in
understanding the cell
cycle and cell cycle
signaling have led to
advances in cancer
treatment
▪Coupled with the ability
to sequence the DNA of
cells in a particular
tumor, treatments are
becoming more
“personalized”
Figure 12.UN02

G1 S
Cytokinesis
Mitosis G2

MITOTIC (M) PHASE

Prophase

Telophase and
Cytokinesis

Prometaphase
Anaphase
Metaphase

© 2017 Pearson Education, Inc.

Figure 12.UN03

© 2017 Pearson Education, Inc.

Figure 12.UN04

© 2017 Pearson Education, Inc.

 Chapter 13

Meiosis and
Sexual Life
Cycles

Lecture Presentations by
Nicole Tunbridge and
© 2017 Pearson Education, Inc.
Kathleen Fitzpatrick
 Variations on a Theme
Offspring resemble their
parents more than they
do unrelated individuals
Heredity is the
transmission of traits
from one generation to
the next
© 2017 Pearson Education, Inc.
 Variations on a Theme
Variation is demonstrated by
the differences in
appearance that offspring
show from parents and
siblings
Genetics is the scientific
study of heredity and
variation
© 2017 Pearson Education, Inc.
Figure 13.1

© 2017 Pearson Education, Inc.

 Concept 13.1: Offspring acquire genes
from parents by inheriting
chromosomes
In a literal sense, children do
not inherit particular
physical traits from their
parents
It is genes that are actually
inherited
© 2017 Pearson Education, Inc.
 Inheritance of Genes
Genes are the units of
heredity and are made up of
segments of DNA
Genes are passed to the
next generation via
reproductive cells called
gametes (sperm and eggs)
© 2017 Pearson Education, Inc.
 Inheritance of Genes
Most DNA is packaged into
chromosomes
Humans have 46 chromosomes in
the nuclei of their somatic cells,
all cells of the body except
gametes and their precursors
A gene’s specific position along a
chromosome is called its locus
© 2017 Pearson Education, Inc.
 Comparison of Asexual and Sexual
Reproduction
In asexual reproduction, a
single individual passes all of
its genes to its offspring without
the fusion of gametes
A clone is a group of genetically
identical individuals from the
same parent
© 2017 Pearson Education, Inc.
 Comparison of Asexual and Sexual
Reproduction
In sexual reproduction, two
parents give rise to offspring
that have unique
combinations of genes
inherited from the two
parents
© 2017 Pearson Education, Inc.
Figure 13.2

0.5 mm

Parent
Bud

(a) Hydra (b) Redwoods

© 2017 Pearson Education, Inc.

Concept 13.2: Fertilization and meiosis
alternate in sexual life cycles
A life cycle is the
generation-to-generation
sequence of stages in
the reproductive history
of an organism
© 2017 Pearson Education, Inc.
 Sets of Chromosomes in Human Cells
Human somatic cells
have 23 pairs of
chromosomes
A karyotype is an
ordered display of the
pairs of chromosomes
from a cell
© 2017 Pearson Education, Inc.
 Sets of Chromosomes in Human Cells
The two chromosomes in each
pair are called homologous
chromosomes, or homologs
Chromosomes in a homologous
pair are the same length and
shape and carry genes
controlling the same inherited
characters
© 2017 Pearson Education, Inc.
Figure 13.3

Application Technique
Pair of homologous
duplicated chromosomes

Centromeres
5 µm

Sister
chromatids
Metaphase
chromosome

© 2017 Pearson Education, Inc.

The sex chromosomes,
which determine the sex
of the individual, are
called X and Y
Human females have a
homologous pair of X
chromosomes (XX)
© 2017 Pearson Education, Inc.
Human males have
one X and one Y
chromosome
The remaining 22 pairs
of chromosomes are
called autosomes
© 2017 Pearson Education, Inc.
Each pair of homologous
chromosomes includes one
chromosome from each
parent
The 46 chromosomes in a
human somatic cell are two
sets of 23: one from the
mother and one from the
father
© 2017 Pearson Education, Inc.
A diploid cell (2n)
has two sets of
chromosomes
For humans, the
diploid number is 46
(2n = 46)
© 2017 Pearson Education, Inc.
In a cell in which DNA
synthesis has occurred,
each chromosome is
replicated
Each replicated
chromosome consists of
two identical sister
chromatids
© 2017 Pearson Education, Inc.
Figure 13.4

Maternal set of chromosomes (n = 3)

2n = 6
Paternal set of chromosomes (n = 3)

Sister chromatids
of one duplicated
chromosome
Centromere

Two nonsister Pair of homologous

chromatids in chromosomes
a homologous pair (one from each set)

© 2017 Pearson Education, Inc.

A gamete (sperm or egg)
contains a single set of
chromosomes and is
thus a haploid cell (n)
For humans, the haploid
number is 23 (n = 23)
© 2017 Pearson Education, Inc.
Each set of 23 consists of 22
autosomes and a single sex
chromosome
In an unfertilized egg (ovum),
the sex chromosome is X
In a sperm cell, the sex
chromosome may be either
X or Y
© 2017 Pearson Education, Inc.
 Behavior of Chromosome Sets in the
Human Life Cycle
Fertilization is the union of gametes
(the sperm and the egg)
The fertilized egg is called a zygote
and has one set of chromosomes
from each parent
The zygote produces somatic cells by
mitosis and develops into an adult
© 2017 Pearson Education, Inc.
At sexual maturity, the
ovaries and testes produce
haploid gametes
Gametes are the only types
of human cells produced
by meiosis, rather than
mitosis
© 2017 Pearson Education, Inc.
Meiosis results in one set
of chromosomes in each
gamete
Fertilization and meiosis
alternate in sexual life
cycles to maintain
chromosome number
© 2017 Pearson Education, Inc.
Figure 13.5
Haploid gametes (n = 23)
Egg (n)

Sperm (n)

MEIOSIS FERTILIZATION

Testis
Ovary
Diploid
zygote
(2n = 46)

Mitosis and
development

Multicellular diploid Haploid (n)

adults (2n = 46) Diploid (2n)
© 2017 Pearson Education, Inc.
 The Variety of Sexual Life Cycles
The alternation of meiosis and
fertilization is common to all
organisms that reproduce
sexually
The three main types of sexual
life cycles differ in the timing
of meiosis and fertilization
© 2017 Pearson Education, Inc.
Figure 13.6

Haploid (n) Haploid multi-

Diploid (2n) cellular organism Haploid unicellular or
(gametophyte) multicellular organism
n Gametes n
n Mitosis n Mitosis Mitosis n Mitosis
n n n
n n
MEIOSIS FERTILIZATION Spores n n
Gametes Gametes n
MEIOSIS FERTILIZATION

2n Zygote 2n MEIOSIS FERTILIZATION

2n
2n Zygote
2n
Mitosis Mitosis Zygote
Diploid Diploid
multicellular multicellular
organism organism
(sporophyte)
(a) Animals (b) Plants and some algae (c) Most fungi and
some protists

© 2017 Pearson Education, Inc.

Gametes are the only haploid
cells in animals
They are produced by meiosis
and undergo no further cell
division before fertilization
Gametes fuse to form a diploid
zygote that divides by mitosis
to develop into a multicellular
organism
© 2017 Pearson Education, Inc.
Figure 13.6a

n Gametes n

MEIOSIS FERTILIZATION

Zygote
2n 2n
Diploid
multicellular Mitosis
organism

Haploid (n)
(a) Animals Diploid (2n)

© 2017 Pearson Education, Inc.

Plants and some algae exhibit an
alternation of generations
This life cycle includes both a
diploid and haploid multicellular
stage
The diploid organism, called the
sporophyte, makes haploid spores
by meiosis

© 2017 Pearson Education, Inc.

Each spore grows by mitosis into a
haploid organism called a
gametophyte
A gametophyte makes haploid
gametes by mitosis
Fertilization of gametes results in a
diploid sporophyte

© 2017 Pearson Education, Inc.

Figure 13.6b
Haploid multi-
cellular organism
(gametophyte)

Mitosis n Mitosis
n n
n n
Spores
Gametes

MEIOSIS FERTILIZATION

2n
2n
Diploid Zygote
multicellular Mitosis
organism
(sporophyte)
Haploid (n)
(b) Plants and some algae Diploid (2n)
© 2017 Pearson Education, Inc.
In most fungi and some protists, the only
diploid stage is the single-celled zygote;
there is no multicellular diploid stage
The zygote produces haploid cells by
meiosis
Each haploid cell grows by mitosis into a
haploid multicellular organism
The haploid adult produces gametes by
mitosis

© 2017 Pearson Education, Inc.

Figure 13.6c

Haploid unicellular or
multicellular organism

Mitosis n Mitosis

n
n n
Gametes n

MEIOSIS FERTILIZATION

2n

Zygote
Haploid (n)
(c) Most fungi and some protists Diploid (2n)
© 2017 Pearson Education, Inc.
Depending on the type of life cycle, either
haploid or diploid cells can divide by
mitosis
However, only diploid cells can undergo
meiosis
In all three life cycles, the halving and
doubling
of chromosomes contributes to genetic
variation
in offspring

© 2017 Pearson Education, Inc.

Concept 13.3: Meiosis reduces the number of
chromosome sets from diploid to haploid
▪ Like mitosis, meiosis is
preceded by the replication of
chromosomes
▪ Meiosis takes place in two
consecutive cell divisions,
called meiosis I and meiosis
II
Concept 13.3: Meiosis reduces the number of
chromosome sets from diploid to haploid
▪ The two cell divisions result in
four daughter cells, rather than
the two daughter cells in mitosis
▪ Each daughter cell has only half
as many chromosomes as the
parent cell
The Stages of Meiosis

▪Chromosomes duplicate
before meiosis
▪The resulting sister
chromatids are closely
associated along their
lengths
The Stages of Meiosis

▪This is called sister

chromatid cohesion
▪The chromatids are
sorted into four haploid
daughter cells
Figure 13.7
Interphase
Pair of
homologous
chromosomes
in diploid
parent cell

Pair of duplicated Chromosomes

homologous duplicate
chromosomes

Sister Diploid cell with

chromatids duplicated
chromosomes
Meiosis I
1
Homologous
chromosomes
separate
Haploid cells with
duplicated chromosomes
Meiosis II
2 Sister chromatids
separate

Haploid cells with unduplicated chromosomes

© 2017 Pearson Education, Inc.
▪Division in meiosis I
occurs in four phases:
▪prophase I
▪metaphase I
▪anaphase I
▪telophase I and
cytokinesis
Prophase I
▪In early prophase I,
each chromosome
pairs with its homolog
and crossing over
occurs
▪X-shaped
regions called
chiasmata are
sites
of crossovers
Metaphase I
▪In metaphase I, pairs
of homologs line up at
the metaphase plate,
with one chromosome
facing each pole
▪Microtubules from one pole
are attached to the
kinetochore of one
chromosome of each pair
▪Microtubules from the other
pole are attached to the
kinetochore of the other
chromosome
Anaphase I
▪In anaphase I, pairs
of homologous
chromosomes
separate
▪One chromosome of
each pair moves
toward opposite
poles, guided by the
spindle apparatus
▪Sister chromatids
remain attached at
the centromere and
move as one unit
toward the pole
Telophase I and
Cytokinesis
▪In the beginning of
telophase I, each half of the
cell has a haploid set of
chromosomes; each
chromosome still consists of
two sister chromatids
▪Cytokinesis usually
occurs
simultaneously,
forming two haploid
daughter cells
▪In animal cells, a
cleavage furrow
forms; in plant
cells, a cell plate
forms
▪No chromosome replication
occurs between the end of
meiosis I and the beginning
of meiosis II because the
chromosomes are already
replicated
Figure 13.8a

MEIOSIS I: Separates homologous chromosomes

Prophase I Metaphase I Anaphase I Telophase I

and Cytokinesis

Centrosome Sister
(with Kinetochore chromatids
centriole (at centromere) remain
Sister pair) attached
chroma- Chiasmata
tids
Spindle Kinetochore
micro- microtubules
tubules

Cleavage
Homologous furrow
Pair of Fragments chromosomes
homo- of nuclear separate
envelope Metaphase
logous plate
chromo-
somes Centromere

© 2017 Pearson Education, Inc.

▪Division in meiosis II also
occurs in four phases:
▪prophase II
▪metaphase II
▪anaphase II
▪telophase II and cytokinesis
▪Meiosis II is very similar to
mitosis
Prophase II
▪In prophase II, a
spindle
apparatus forms
▪In late prophase II,
chromosomes (each
still composed of two
chromatids) move
toward the metaphase
plate
Metaphase II
▪In metaphase II, the
sister chromatids are
arranged at the
metaphase plate
▪Because of crossing
over in meiosis I, the
two sister chromatids
of each chromosome
are no longer
genetically identical
▪The kinetochores of
sister chromatids
attach to
microtubules
extending from
opposite poles
Anaphase II
▪In anaphase II,
the sister
chromatids
separate
▪The sister chromatids
of each chromosome
now move as two
newly individual
chromosomes toward
opposite poles
Telophase II and
Cytokinesis
▪In telophase II, the
chromosomes arrive
at opposite poles
▪Nuclei form, and
the chromosomes
begin
decondensing
▪Cytokinesis
separates the
cytoplasm
▪At the end of
meiosis, there are
four daughter cells,
each with a haploid
set of unreplicated
chromosomes
▪Each daughter
cell is genetically
distinct from the
others and from
the parent cell
Figure 13.8b

MEIOSIS II: Separates sister chromatids

Prophase II Metaphase II Anaphase II Telophase II

and Cytokinesis

Sister chromatids
separate Haploid daughter
cells forming

© 2017 Pearson Education, Inc.

Crossing Over and Synapsis During Prophase I

▪After interphase, the sister

chromatids are held
together by proteins called
cohesins
▪The nonsister chromatids
are broken at precisely
corresponding positions
Crossing Over and Synapsis During Prophase I
▪ A zipper-like structure called the
synaptonemal complex holds the
homologs together tightly
▪ During synapsis, DNA breaks are
repaired, joining DNA from one
nonsister chromatid to the
corresponding segment of another
Figure 13.9

Pair of homologous Sister Synaptonemal

chromosomes: chromatids complex
DNA Centromere DNA Sister
Paternal chromatids
breaks breaks sister
Cohesins
chromatids

Maternal
sister Crossovers
1 chromatids 3

Synaptonemal
Chiasmata
complex forming

2 4

© 2017 Pearson Education, Inc.

A Comparison of Mitosis and Meiosis

▪Mitosis conserves the

number of chromosome
sets, producing cells that
are genetically identical
to the parent cell
A Comparison of Mitosis and Meiosis

▪Meiosis reduces the number

of chromosomes sets from
two (diploid) to one
(haploid), producing cells
that differ genetically from
each other and from the
parent cell
▪Three events are unique to
meiosis, and all three occur
in meiosis I
▪Synapsis and crossing over in
prophase I: Homologous
chromosomes physically
connect and exchange
genetic information
▪Homologous pairs at
the metaphase plate
▪Separation of
homologs during
anaphase I
Figure 13.10a

MITOSIS MEIOSIS
Parent cell Chiasma MEIOSIS I

Prophase Prophase I
Chromosome Chromosome
Duplicated Pair of
duplication duplication
chromosome 2n = 6 duplicated
homologs
Individual Pairs of
Metaphase homologous Metaphase I
chromosomes
line up. chromosomes
line up.
Anaphase Anaphase I
Sister chromatids Homologs
Telophase separate. Telophase I
separate.

Daughter
Sister cells of
meiosis I
chromatids MEIOSIS II
2n 2n separate.
Daughter cells n n n n
of mitosis Daughter cells of meiosis II

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▪Sister chromatid cohesion
allows sister chromatids to
stay together through
meiosis I
▪In mitosis, cohesins are
cleaved at the end of
metaphase
▪In meiosis, cohesins are
cleaved along the
chromosome arms in
anaphase I (separation of
homologs) and at the
centromeres in anaphase II
(separation of sister
chromatids)
Concept 13.4: Genetic variation produced in
sexual life cycles contributes to evolution
▪ Mutations (changes in an organism’s
DNA) are the original source of
genetic diversity
▪ Mutations create different versions of
genes called alleles
▪ Reshuffling of alleles during sexual
reproduction produces genetic
variation
Concept 13.4: Genetic variation produced in
sexual life cycles contributes to evolution

▪Mutations (changes
in an organism’s
DNA) are the original
source of genetic
diversity
Concept 13.4: Genetic variation produced in
sexual life cycles contributes to evolution

▪Mutations create different

versions of genes called
alleles
▪Reshuffling of alleles during
sexual reproduction
produces genetic variation
Origins of Genetic Variation Among Offspring

▪The behavior of
chromosomes during
meiosis and fertilization
is responsible for most of
the variation that arises
in each generation
Origins of Genetic Variation Among Offspring

▪Three mechanisms
contribute to genetic
variation:
▪Independent assortment of
chromosomes
▪Crossing over
▪Random fertilization
Independent Assortment of Chromosomes

▪Homologous pairs
of chromosomes
orient randomly at
metaphase I of
meiosis
Independent Assortment of Chromosomes

▪In independent assortment,

each pair of chromosomes
sorts maternal and paternal
homologs into daughter
cells independently of the
other pairs
▪The number of
combinations possible
when chromosomes
assort independently into
n
gametes is 2 , where n is
the haploid number
▪For humans (n =
23), there are more
23
than 8 million (2 )
possible
combinations of
chromosomes
Figure 13.11_1

Possibility 1 Possibility 2

Two equally probable

arrangements of
chromosomes at
metaphase I

© 2017 Pearson Education, Inc.

Figure 13.11_2

Possibility 1 Possibility 2

Two equally probable

arrangements of
chromosomes at
metaphase I

Metaphase II

© 2017 Pearson Education, Inc.

Figure 13.11_3

Possibility 1 Possibility 2

Two equally probable

arrangements of
chromosomes at
metaphase I

Metaphase II

Daughter
cells
Combination 1 Combination 2 Combination 3 Combination 4

© 2017 Pearson Education, Inc.

Crossing Over

▪Crossing over
produces
recombinant
chromosomes, which
combine DNA inherited
from each parent
Crossing Over

▪Crossing over
contributes to genetic
variation by combining
DNA from two parents
into a single
chromosome
Crossing Over

▪In humans, an average

of one to three
crossover events
occurs per
chromosome
Figure 13.12_1
Prophase I Nonsister chromatids
of meiosis held together
during synapsis
Pair of
homologs 1 Synapsis and
crossing over
Chiasma

Centromere

TEM

© 2017 Pearson Education, Inc.

Figure 13.12_2
Prophase I Nonsister chromatids
of meiosis held together
during synapsis
Pair of
homologs 1 Synapsis and
crossing over
Chiasma
2 Movement to
the metaphase I
plate
Centromere

TEM
Anaphase I

© 2017 Pearson Education, Inc.

Figure 13.12_3
Prophase I Nonsister chromatids
of meiosis held together
during synapsis
Pair of
homologs 1 Synapsis and
crossing over
Chiasma
2 Movement to
the metaphase I
plate
Centromere
3 Breakdown of
TEM
Anaphase I proteins holding
sister chromatid
arms together

Anaphase II

© 2017 Pearson Education, Inc.

Figure 13.12_4
Prophase I Nonsister chromatids
of meiosis held together
during synapsis
Pair of
homologs 1 Synapsis and
crossing over
Chiasma
2 Movement to
the metaphase I
plate
Centromere
3 Breakdown of
TEM
Anaphase I proteins holding
sister chromatid
arms together

Anaphase II

Daughter
cells
Recombinant chromosomes
© 2017 Pearson Education, Inc.
Random Fertilization

▪Random fertilization
adds to genetic
variation because any
sperm can fuse with
any ovum (unfertilized
egg)
Random Fertilization
▪ The fusion of two gametes
(each with 8.4 million possible
chromosome combinations
from independent assortment)
produces a zygote with any of
about 70 trillion diploid
combinations
▪Crossing over adds
even more variation
▪Each zygote has a
unique genetic
identity
The Evolutionary Significance of Genetic
Variation Within Populations
▪ Natural selection results in the
accumulation of genetic variations
favored by the environment
▪ Sexual reproduction contributes to
the genetic variation in a
population, which originates from
mutations
The Evolutionary Significance of Genetic
Variation Within Populations

▪ Animals that always reproduce

asexually are quite rare
▪ Organisms like the bdelloid
rotifer increase their genetic
diversity through incorporation
of foreign DNA from the
environment
Figure 13.UN03

F
H

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