MI TOSIS

MI T
Spore OS
IS
S IS (n) Developing haploid
IO
organism (n)
Spores are
made by
meiosis
Sperm (n)
Mature
haploid
organism (n)
M
IT Zygote
OS Sperm
IS (2n) swim to egg
Mature diploid Egg (n)
organism (2n)
FERT ILI ZAT ION

Chapter 13
MEIOSIS

In this chapter you will learn how

Meiosis promotes genetic diversity and
allows the benefits of sex
starting with by examining by asking

How does meiosis How meiosis produces Why does meiosis

occur? genetic variation exist?
13.1 13.4
looking at
and comparing to Purifying-selection
Independent assortment, hypothesis
Mitosis
crossing over, and
Ch. 12 fertilization
13.2 Changing-environment
hypothesis
then asking
What happens when
things go wrong?
13.3

1
 Learning Objectives
o Define homologous, autosomes, and sex
chromosomes, allele, ploidy (haploid, diploid,
polyploid and aneuploid), and sexual and asexual
reproduction.

o Compare the different purposes of mitosis and

meiosis.

o Explain the leading hypothesis for why meiosis

occurs (changing environment hypothesis) even
though asexual reproduction is more efficient at
producing new offspring.

Learning Objectives
o Describe what happens in meiosis I & II.

o Describe how meiosis mixes up the genes on each

chromosome. Recall when in meiosis homologous
chromosomes pair up and when synapsis (crossing
over) occurs.

o Explain how mistakes in meiosis, nondisjunction,

lead to cells with the wrong number of
chromosomes and explain why this leads to
improper development.

2
 Introduction
v During sexual reproduction, a sperm and an egg
unite to form a new individual.
v This process is called fertilization.

v Meiosis is nuclear division that halves the number

of chromosomes.

v Meiosis precedes the formation of gametes (egg

and sperm) in some organisms including animals.

Drosophila chromosomes and photographed through

drawn by Nettie Stevens a modern microscope.
in 1908

2 3
2 3

4 4
X Y
Autosomes Sex
(24) X Y
chromosomes

3
 Chromosomes Come in
Distinct Types
v Each organism has a characteristic number of
chromosomes.

v The karyotype is the number and types of

chromosomes present in an organism.

v Sex chromosomes determine the sex of the individual

v All other chromosomes are autosomes.

v Humans have 46 chromosomes in every cell except their

gametes.
v 1 pair of sex chromosomes.
v 22 pairs of autosomes.

4
Homologous Chromosomes
Have the Same Genes
v A gene is a section of DNA that influences one or more
hereditary traits in an individual.
v Different versions of a specific gene are called alleles.
v Homologs (homologous chromosomes) carry the
same genes in
Homologous
the same chromosomes
locations, but (replicated)
each one may
contain Gene for eye color Gene for eye color
(allele for red eyes) (allele for purple eyes)
different alleles.

Drosophila autosome 2

The Concept of Ploidy

v The haploid number n indicates the number of
distinct types of chromosomes present.

v A cells ploidy (n, 2n, 3n, etc.) indicates the

number of each type of chromosome present.
Organisms are called
v haploid (n) if cells contain just one of each type of
chromosome.
v diploid (2n) if cells contain two versions of each type of
chromosome.
v Diploid cells have one paternal chromosome and one
maternal chromosome.
v polyploid (3n, 4n, etc.) if cells contain three or more
versions of each type of chromosome.

5
6
 An Overview of Meiosis
v Meiosis reduces chromosome number by
half. In diploid organisms, the products of
meiosis are haploid.

v Just before meiosis begins, each

chromosome in the diploid (2n) parent cell is
replicated.
v When replication is complete, each chromosome
consists of two identical sister chromatids
attached at the centromere.

Maternal Paternal
chromosome chromosome
Nuclear envelope

Replication

Replicated Centromere
chromosome
Sister chromatids

Homologous pair of
replicated chromosomes

7
 An Overview of Meiosis
v Meiosis consists of two cell divisions
1. Meiosis I
2. Meiosis II

v The two divisions occur consecutively but differ

sharply

An Overview of Meiosis I
v During meiosis I, the diploid (2n) parent cell
produces two haploid (n) daughter cells.

v The homologs in each chromosome pair

separate and go to different daughter cells.

v Although the daughter cells are haploid (n),

each chromosome still consists of two
identical sister chromatids.

8
 An Overview of Meiosis II
v During meiosis II, the sister chromatids of
each chromosome separate and go to
different daughter cells.

v The four haploid daughter cells produced by

meiosis II also have one of each type of
chromosome, but now the chromosomes are
unreplicated.

Parent cell is
diploid (2n)
and contains
MEIOSIS I

a homologous
pair of
replicated
chromosomes
Homologs
separate Daughter
cells are
haploid
(n) and
MEIOSIS II

Sister contain
chromatids just one
separate homolog

Four daughter cells contain one chromosome each (n),

In animals, these cells become gametes.

9
 Meiosis is a Reduction
Division
v The outcome of meiosis is a reduction in
chromosome number. For this reason,
meiosis is known as a reduction division.

v In most plants and animals, the original cell

is diploid and the four daughter cells are
haploid.
v In animals, these daughter cells become
gametes via a process called gametogenesis.

Fertilization Results in a
Diploid Zygote
v When two haploid gametes fuse during
fertilization, a full complement of chromosomes is
restored. The cell that results from fertilization is
diploid and is called a zygote.

v In this way, each diploid individual receives a

haploid chromosome set from both its mother and
its father.
v Homologs are therefore referred to as being either
maternal chromosomes, from the mother, or
paternal chromosomes, from the father.

10
 Female Male
gamete gamete
(egg) (sperm)
(n) (n)
Fertilization

Diploid offspring
contains homologous
pair of chromosomes

Zygote (2n)

The Life Cycle of a

Sexual Organism
v An animals life cycle summarizes life from
fertilization through offspring production.

v Meiosis in an adult produces haploid gametes that

combine during fertilization to form a diploid zygote,
which develops, through mitosis, into an adult of the
next generation.

11
 Diploid (2n) MEIOSIS
Haploid (n)

Amount of hereditary
material reduced by half

Haploid
Diploid gametes (n)
Sperm Egg
adult
(2n)

ON
Normal amount

TI
of hereditary

ZA
material restored

LI
R TI
FE
(dur M
ing ITOSIS Zygote
dev
elop (2n)
men
t)

The Phases of Meiosis I

v Meiosis I is a continuous process with five
distinct phases:
1. Early prophase I
2. Late prophase I
3. Metaphase I
4. Anaphase I
5. Telophase I

12
 The Phases of Meiosis I
v Early Prophase I: The homolog pairs come together
in a pairing process called synapsis. The structure
that results from synapsis is called a tetrad,
consisting of two homologs. The chromatids of the
homologs are called non-sister chromatids.

v Late Prophase I: These non-sister chromatids begin

to separate. Exchange or crossing over between
homologous non-sister chromatids occurs where
chiasmata are formed during this stage.

The Phases of Meiosis I

v Metaphase I: The tetrads line up at the metaphase
plate.

v Anaphase I: The paired homologs separate and

begin to migrate to opposite ends of the cell.

v Telophase I: The homologs finish migrating to the

poles of the cell.

v Then the cell

divides in the
process of
cytokinesis.

13
Nuclear Bivalent (4 chromatids from
envelope 2 homologous chromosomes)
Replicated Non-sister Spindle
chromosomes chromatids apparatus Chiasma

2n Maternal chromosomes
In this example,
n = 3 so 2n = 6 Paternal chromosomes

1. Interphase 2. Early prophase I 3. Late prophase I 4. Metaphase I 5. Anaphase I

The Result of Meiosis I

v The end result of meiosis I is that one
chromosome of each homologous pair is
distributed to a different daughter cell.

v A reduction division has occurred.

v The daughter cells of meiosis I are haploid and
are still in the form of sister chromatids.

v Crossing over.

14
 The Phases of Meiosis II
v Like meiosis I, meiosis II is a continuous
process, but with four distinct phases:
1. Prophase II
2. Metaphase II
3. Anaphase II
4. Telophase II

n n

6. Telophase I 7. Prophase II 8. Metaphase II 9. Anaphase II 10. Telophase II

and Cytokinesis and Cytokinesis

15
 The Phases of Meiosis II
v Prophase II: The spindle apparatus forms and one
spindle fiber attaches to the centromere of each sister
chromatid.

v Metaphase II: Replicated chromosomes line up at the

metaphase plate.

v Anaphase II: Sister chromatids separate. The resulting

daughter chromosomes begin moving to opposite sides
of the cell.

v Telophase II: Chromosomes arrive at opposite sides of

the cell. A nuclear envelope forms around each haploid
set of chromosomes.

v Each cell undergoes cytokinesis.

The Result of Meiosis II

v Meiosis II results in four haploid cells, each
with one of each type of chromosome.
v Thus, one diploid cell with replicated
chromosomes gives rise to four haploid
cells with unreplicated chromosomes.

16
 Order these cells!

Comparison of
Meiosis & Mitosis
v The key difference between the two processes is
that homologs pair in meiosis, but do not in mitosis.

v Because homologs pair in prophase of meiosis I,

they can migrate to the metaphase plate together
and then separate during anaphase of meiosis I,
resulting in a reduction division.

v Meiosis thus produces four daughter cells with half

the genetic material of the parents.

v Mitosis produces two daughter cells that are

genetically identical to the parent cells.

17
 MITOSIS MEIOSIS

2n Diploid parent cell 2n Diploid parent cell

Chromosome replication Chromosome replication

2n Prophase 2n Prophase I

2n Metaphase 2n Metaphase I

Anaphase and
Telophase

2n n Anaphase I and
Telophase I

Two diploid daughter

cells of mitosis Meiosis II

Four haploid daughter cells of meiosis

Where Does Crossing

Over Occur?
v After replication, sister chromatids stay
tightly joined along their entire length.

v When homologs synapse, two pairs of non-

sister chromatids are brought close together
and are held there by a network of proteins
called the synaptonemal complex.

v Crossing over occurs when chromosomal

segments are swapped between adjacent
homologs.

18
 A Closer Look at Crossing
Over
v At each point where crossing over occurs, the non-
sister chromatids from each homolog get physically
broken at the same point and attached to each
other. As a result, segments of maternal and
paternal chromosomes are swapped.

v Crossing over can occur at many locations along

the length of such synapsed homologs. Both sets of
non-sister chromatids may undergo crossing over,
resulting in the swapping of segments between
maternal and paternal chromosomes.

Bivalent Crossover between

non-sister chromatids

Break in
chromatid
Cohesin DNA Chiasma
proteins
Kinetochore
microtubule

Kinetochores

Homologs

Synaptonemal
complex
Sister chromatids
1. Condensation 2. Pairing 3. Synapsis (bivalent 4. Partial separation
formation) of homologs

19
 The Consequences of
Meiosis
v Independent shuffling of maternal and
paternal chromosomes and crossing over
during meiosis I result in four gametes with a
chromosome composition different from that
of the parent cells.

v The changes in chromosomes produced by

meiosis and fertilization are significant
because chromosomes contain the cells
hereditary material.

Sexual Reproduction Leads to

Greater Variation
v Offspring produced during asexual reproduction
are clones that are genetically identical to one
another as well as to the parent.

v In contrast, offspring produced by sexual

reproduction, the fusion of gametes, have a
chromosome makeup different from that of one
another and from that of either parent.

v Genetic variation in sexual reproduction results from

independent assortment, crossing over and random
combination of gametes.

20
 Independent Assortment
Produces Genetic Variation
v Separation and distribution of homologous
chromosomes during meiosis I can result in a
variety of combinations of maternal and paternal
chromosomes.

v Each daughter cell gets a random assortment of

maternal and paternal chromosomes, and thus
genes, which generates a great deal of genetic
diversity in the subsequent gametes.

v Humans, with a haploid chromosome number of 23,

can produce 223 (~ 8.4 million) different
combinations of chromosomes in gametes.

Example: Individual with different alleles of two genes

Maternal Paternal Maternal Paternal

chromosome chromosome chromosome chromosome

Allele that Allele that

contributes contributes
to black to red hair
hair
Allele that Allele that A hair-color gene
contributes contributes
to brown to green eyes
eyes
An eye-color gene

21
During meiosis I, bivalents can line up two different
ways before the homologs separate.

OR

Brown eyes Green eyes Brown eyes Green eyes

Black hair Red hair Red hair Black hair

The Role of Crossing Over

v Crossing over produces new combinations
of alleles on the same chromosome,
combinations that did not exist in each
parent.

v Crossing over is a form of genetic

recombination that increases the genetic
variability of gametes produced by meiosis
beyond that produced by random
assortment of chromosomes.

22
How Does Fertilization Affect
Genetic Variation?
v Crossing over and the random mixing of
maternal and paternal chromosomes ensure
that each gamete is genetically unique.

v The genetic variation introduced during

meiosis ensures that even in self-
fertilization, where gametes from the same
individual combine, the offspring will be
genetically different from the parent.

The red and blue

chromosomes can line
up in different ways
during metaphase

OR

2n = 4
2. Crossing over
1. Parent cell
3. Independent
assortment

4. Gametes 5. Fertilization

23
Outcrossing Further Increases
Genetic Variation
v In many sexually reproducing species, gametes from
different individuals combine to form offspring, a
process called outcrossing.

v Outcrossing increases the genetic diversity of the

offspring because chromosomes from two different
parents are combined.
v In humans, this means that two parents can
potentially produce 223 combinations of gametes
v With fertilization = 8.4 million x 8.4 million
= 70.6 x 1012 genetically distinct offspring.
This does not even take additional variation from
crossing over into consideration!

Evolutionary
Consequences of Sex
v Sexual reproduction increases genetic diversity
through three key mechanisms

1. Crossing over
2. Independent
assortment
3. Outcrossing

24
 Why Does Meiosis Exist?
v Sexual reproduction is common among
multicellular organisms, but organisms in
most lineages of the tree of life undergo
asexual reproduction

v So why reproduce by fusing gametes?

The Paradox of Sex

v The mathematical model of John Maynard Smith
predicts that asexually reproducing organisms should
reproduce faster and outcompete similar organisms
that invest in sexual reproduction.

v Asexual reproduction is much more efficient than

sexual reproduction because no males are produced.
Asexual reproduction Sexual reproduction

Generation 1

There are only half as

many child-producing
offspring in the sexual
population as in the
Generation 2 asexual population

Generation 3

25
 The Purifying Selection
Hypothesis
v In asexual reproduction, a damaged gene will
be inherited by all of that individuals offspring.

v Sexually reproducing individuals are likely to

have offspring that lack deleterious alleles
present in the parent.

v Natural selection against deleterious alleles is

called purifying selection. Over time, purifying
selection should steadily reduce the numerical
advantage of asexual reproduction.

The Changing
Environment Hypothesis
v Offspring produced by sexual reproduction
are genetically different from their parents
v More likely to survive and produce offspring if
the environment changes

v Offspring produced by asexual reproduction

are genetically identical to their parents
v Less likely to survive and produce offspring if the
environment changes.

26
 Testing the Changing
Environment Hypothesis
v If a new strain of disease-causing agent evolves,
then all of the asexually produced offspring are
likely to be susceptible to that new strain.

v But if the offspring are genetically varied, then it is

likely that at least some offspring will have
combinations of alleles that enable them to fight off
the new disease and produce offspring of their own.

v Studies support the changing environment

hypothesis. Sexual reproduction may be an
adaptation that increases the fitness of individuals in
certain environments.

27
28
 Does exposure to evolving pathogens favor outcrossing?
In environments where evolving pathogens are present, sexual reproduction
by outcrossing will be favored.
The presence of evolving pathogens will not favor outcrossing.

1. Start with a pathogen free

population of roundworms.

2. Divide the population;

grow subgroups in
absence and presence
Grow without Grow with
of evolving pathogen.
pathogen pathogen

3. Assess the rate of

outcrossing over many
generations.

The rate of outcrossing will increase in response to exposure by a pathogen.

The rate of outcrossing will not be influenced by a pathogen.

With
pathogen
Outcrossing rate

Without pathogen

Generation
Exposure to evolving pathogens favors outcrossing.

29
 Mistakes in Meiosis
v If a mistake occurs during meiosis and

v the chromosomes from the parent cells are

not properly distributed to each daughter cell

v the resulting gametes will contain an

abnormal set of chromosomes.

How Do Mistakes Occur?

v For a gamete to get one complete set of
chromosomes, each pair of homologous
chromosomes must separate from each other
during the first meiotic division, and the sister
chromatids must separate from each other and
move to opposite poles during meiosis II.

v If both homologs or both sister chromatids move to

the same pole of the parent cell, the products of
meiosis will be abnormal. This sort of meiotic error
is referred to as nondisjunction.

30
 n+1

n+1

n-1

2n = 4
n=2 n-1

1. Meiosis I starts 2. Nondisjunction 3. Meiosis II occurs 4. Aneuploidy results.

normally. Bivalents occurs with one normally. All gametes have too
line up in middle of set of homologs. many or too few
cell. chromosomes.

Types of Nondisjunction
v If nondisjunction occurs in meiosis I, two gametes
will have an extra copy of a chromosome (causing a
condition called trisomy), and two gametes will lack
that chromosome (monosomy).
v An example of trisomy is Down syndrome, which is
caused by an extra copy of chromosome 21.

v If nondisjunction occurs in meiosis II, two gametes

will be normal, one cell will have an extra copy of a
chromosome, and one gamete will lack that
chromosome.

31
 Frequency of
Nondisjunction
v Nondisjunction may occur in as many 10 percent of
meiotic divisions.

v Aneuploid zygotes (those with too few or too many

chromosomes) typically do not survive to produce
viable offspring.

v Mistakes in meiosis are the leading cause of

spontaneous abortion (miscarriage) in humans.

Why Do Mistakes Occur?

v Meiotic errors appear to be accidental, with
no genetic predisposition.

v Maternal age
Incidence of Down syndrome

is an
per number of births

important
factor in the
frequency of
trisomy 21.

Age of mother (years)

32
 Most Common Human
Aneuploidy Disorders
v Aneuploidy of chromosome 21 leads to
Down Syndrome.

v Sex chromosome aneuploidy can occur in

many different forms:
v Klinefelter syndrome develops in XXY males.
v Trisomy X (karyotype XXX).
v Females with Turner Syndrome have
monosomy their karyotype is XO (they are
lacking a second X chromosome) and are
usually sterile.

33