Essential idea: Alleles segregate during meiosis allowing new

3.3 Meiosis combinations to be formed by the fusion of gametes.

The micrographs above show the formation of bivalents (left) and the segregation caused by
both anaphase I and II (right). These possesses combined with crossing over and random
orientation ensure a near infinite variation of genetic information between gametes.

By Chris Paine
https://bioknowledgy.weebly.com/
https://s10.lite.msu.edu/res/msu/botonl/b_online/e09/meiosea.htm
10.1 Meiosis (AHL)

Essential idea: Meiosis leads to independent assortment of

chromosomes and unique composition of alleles in daughter cells.

The family portrait shows large amounts of variation within a family despite sharing
a lot of genes. This shows the potential of crossing over and independent
assortment to create near infinite variation in gametes and hence in offspring too.

By Chris Paine
https://bioknowledgy.weebly.com/
https://upload.wikimedia.org/wikipedia/commons/9/95/Coloured-family.jpg
Understandings
Statement Guidance
3.3.U1 One diploid nucleus divides by meiosis to produce four
haploid nuclei.
3.3.U2 The halving of the chromosome number allows a
sexual life cycle with fusion of gametes.
3.3.U3 DNA is replicated before meiosis so that all
chromosomes consist of two sister chromatids.
3.3.U4 The early stages of meiosis involve pairing of The process of chiasmata formation need not be
homologous chromosomes and crossing over followed explained.
by condensation.
3.3.U5 Orientation of pairs of homologous chromosomes prior
to separation is random.
3.3.U6 Separation of pairs of homologous chromosomes in the
first division of meiosis halves the chromosome
number.
3.3.U7 Crossing over and random orientation promotes
genetic variation.
3.3.U8 Fusion of gametes from different parents promotes
genetic variation.
Applications and Skills
Statement Guidance
3.3.A1 Non-disjunction can cause Down syndrome and other
chromosome abnormalities.
3.3.A2 Studies showing age of parents influences chances of
non-disjunction.
3.3.A3 Description of methods used to obtain cells for
karyotype analysis e.g. chorionic villus sampling and
amniocentesis and the associated risks.
3.3.S1 Drawing diagrams to show the stages of meiosis Drawings of the stages of meiosis do not need to
resulting in the formation of four haploid cells. include chiasmata. Preparation of microscope
slides showing meiosis is challenging and
permanent slides should be available in case no
cells in meiosis are visible in temporary mounts.
Understandings, Applications and Skills
Statement Guidance
10.1.U1 Chromosomes replicate in interphase before meiosis.
10.1.U2 Crossing over is the exchange of DNA material between
non-sister homologous chromatids.
10.1.U3 Crossing over produces new combinations of alleles on
the chromosomes of the haploid cells.
10.1.U4 Chiasmata formation between non-sister chromatids
can result in an exchange of alleles.
10.1.U5 Homologous chromosomes separate in meiosis I.
10.1.U6 Sister chromatids separate in meiosis II.
10.1.U7 Independent assortment of genes is due to the random
orientation of pairs of homologous chromosomes in
meiosis I.
10.1.S1 Drawing diagrams to show chiasmata formed by Diagrams of chiasmata should show sister
crossing over. chromatids still closely aligned, except at the point
where crossing over occurred and a chiasma was
formed.
Review: 1.6.U1 Mitosis is division of the nucleus into two genetically identical daughter nuclei.

People
Meet
And
Talk
http://www.flickr.com/photos/chuckp/252924532/
Review: 1.6.U1 Mitosis is division of the nucleus into two genetically identical daughter nuclei.

Prophase
Metaphase
Anaphase
Telophase
http://www.flickr.com/photos/chuckp/252924532/
Review: 1.6.U1 Mitosis is division of the nucleus into two genetically identical daughter nuclei.

Use the animated tutorials to learn about mitosis

http://highered.mheducation.com/sites/0072495
855/student_view0/chapter2/animation__mitosis http://www.sumanasinc.com/webcontent/animations/content
_and_cytokinesis.html /mitosis.html

http://www.johnkyrk.com/mitosis.html http://outreach.mcb.harvard.edu/animations/cellcycle.
swf
Nature of Science: Making careful observations—meiosis was discovered by microscope examination of
dividing germ-line cells. (1.8)

1876 - German biologist Oscar Hertwig recognized the role of the cell nucleus during
inheritance and chromosome reduction during meiosis from work on Sea Urchins.

1883 - Belgian zoologist Edouard Van Beneden discovered in the roundworm

Ascaris how chromosomes organized meiosis (the production of gametes).

1890 - The significance of meiosis for reproduction and inheritance was first
described by German biologist August Weismann who noted that two cell
divisions were necessary to transform one diploid cell into four haploid cells.

In the 19th century was very difficult to observe the behaviour of

chromosomes in cell: the choice of organism and tissue, slide
preparation and interpreting microscope images are all difficult to
do successfully. It therefore it took years of careful examination by
Scientists to discover and fully understand meiosis.

http://www.ijdb.ehu.es/web/paper.php?doi=1627480
3.3.U1 One diploid nucleus divides by meiosis to produce four haploid nuclei.

Meosis is a reduction division of the nucleus to form haploid gametes

One diploid (2N) body cells contain a homologous Chromosomes are replicated to
pair of each chromosome (except for sex cells) form sister chromatids

Four haploid (N) gametes contain

one of each chromosome

First
division
of the
nucleus

Second division of the nucleus

Edited from: https://commons.wikimedia.org/wiki/File:Diagram_of_meiosis.svg
3.3.U1 One diploid nucleus divides by meiosis to produce four haploid nuclei.

Meosis is a reduction division of the nucleus to form haploid gametes

The animations
are a great way
to visualise the
http://www.sumanasinc.com/webcontent/animations/c
ontent/meiosis.html
process –
watch and take
notes.
http://www.biostudio.com/d_%20Meiosis.htm

http://highered.mheducation.com/sites/dl http://www.stolaf.edu/people/giannini/fla
/free/0072437316/120074/bio19.swf shanimat/celldivision/meiosis.swf
Outline the differences between the behaviour of
chromosomes in Mitosis and Meiosis
5 marks
Outline the differences between the behaviour of
chromosomes in Mitosis and Meiosis
5 marks

Mitosis Meiosis

One division Two divisions

Diploid cells produced Haploid gametes produced

No crossing-over in prophase Crossing-over in prophase I

No chiasmata formation Chiasmata form

Homologous pairs do not associate and Homologous pairs associate as bivalents

line up at the equator in metaphase and lined up at the equator in metaphase I
Homologous pairs separate in anaphase I
Sister chromatids separate in anaphase
Sister chromatids separate in anaphase II
Review: 3.3.U3 DNA is replicated before meiosis so that all chromosomes consist of two sister chromatids.

Interphase
In the S-phase of the interphase
before meiosis begins, DNA replication
takes place.

Chromosomes are replicated and

these copies are attached to each
other at the centromere.

The attached chromosome and its

copy are known as sister chromatids.

Following S-phase, further growth and

preparation take place for meiosis.
10.1.U1 Chromosomes replicate in interphase before meiosis.

An homologous pair of chromosomes…

10.1.U1 Chromosomes replicate in interphase before meiosis.

An homologous pair of chromosomes…

…replicates during S-phase of interphase…
10.1.U1 Chromosomes replicate in interphase before meiosis.

An homologous pair of chromosomes…

…replicates during S-phase of interphase…

centromere

sister chromatids

…giving two pairs of sister chromatids,

each joined at the centromere.
Review: 3.3.S1 Drawing diagrams to show the stages of meiosis resulting in the formation of four haploid cells. AND 3.3.U4
The early stages of meiosis involve pairing of homologous chromosomes and crossing over followed by condensation.

Prophase I
The homologous
chromosomes
associate with each
other to form
bivalents (synapsis).

DNA supercoils and chromosomes

condense

nuclear membrane dissolves

centrioles migrate to the

poles of the cell.

Crossing-over between non-sister

chromatids can take place. This results
in recombination of alleles and is a
source of genetic variation in gametes.
Edited from: http://www.slideshare.net/gurustip/meiosis-ahl http://www2.sunysuffolk.edu/gambier/micrographs/lateprophase3.htm
10.1.U2 Crossing over is the exchange of DNA material between non-sister homologous chromatids. AND
10.1.U4 Chiasmata formation between non-sister chromatids can result in an exchange of alleles.

The homologous pair associates during prophase I,

through synapsis…

…making a bivalent.
10.1.U2 Crossing over is the exchange of DNA material between non-sister homologous chromatids. AND
10.1.U4 Chiasmata formation between non-sister chromatids can result in an exchange of alleles.

Crossing-over might take place between non-sister

chromatids in prophase I…

The point of crossing-over

between non-sister
chromatids is called the
chiasma (pl. chiasmata).

…leading to recombination of alleles.

10.1.U2 Crossing over is the exchange of DNA material between non-sister homologous chromatids. AND
10.1.U4 Chiasmata formation between non-sister chromatids can result in an exchange of alleles.

The formation of chiasma is common and

is thought to be essential for meiosis Inchiasmata
humans the average number of
per bivalent (tetrad) is just
over two.

How many chiasma can you identify in the micrographs?

a. b.

Chiasma occur between the sister chromatids and therefore should be appear the the
central space.

http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/C/Chiasmata013.jpg
http://nzetc.victoria.ac.nz/etexts/Bio13Tuat01/Bio13Tuat01_063a(h280).jpg
10.1.U2 Crossing over is the exchange of DNA material between non-sister homologous chromatids. AND
10.1.U4 Chiasmata formation between non-sister chromatids can result in an exchange of alleles.

The formation of chiasma is common and

is thought to be essential for meiosis Inchiasmata
humans the average number of
per bivalent (tetrad) is just
over two.

How many chiasma can you identify in the micrographs?

a. b.
3 4
2
2 centromere 1 5

Chiasma occur between the sister chromatids and therefore should be appear the the
central space.

http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/C/Chiasmata013.jpg
http://nzetc.victoria.ac.nz/etexts/Bio13Tuat01/Bio13Tuat01_063a(h280).jpg
10.1.U3 Crossing over produces new combinations of alleles on the chromosomes of the haploid cells.

Crossing-over
Increases genetic variation through
recombination of linked alleles.

Synapsis Chiasma Formation Recombination

Homologous chromosomes Neighbouring non-sister chromatids As a result, alleles are
associate are cut at the same point. swapped between non-
A Holliday junction forms as the DNA sister chromatids.
of the cut sections attach to the
open end of the opposite non-sister
chromatid.
10.1.U3 Crossing over produces new combinations of alleles on the chromosomes of the haploid cells.

Crossing-Over
Increases genetic variation through
recombination of linked alleles.

Crossing over leads to more variation in gametes.

This is the standard notation for writing genotypes of alleles on linked

genes (more of this later when we study 10.2 Inheritance AHL)
Edited from: http://www.slideshare.net/gurustip/meiosis-ahl
10.1.S1 Drawing diagrams to show chiasmata formed by crossing over.

Crossing-Over Drawing clear diagrams helps when

communicating with others
Sister Un-recombined
centromere
chromatids chromatids

chiasma
Homologous
Recombined
pair
chromatids
Key points when drawing chiasmata:
• Use colour of shading to indicate the each of the sister chromatids
• Remember the homologous chromosomes are in synapsis – before and
after crossing over the chromosomes should be shown close together.
Review: 3.3.S1 Drawing diagrams to show the stages of meiosis resulting in the formation of four haploid
cells. AND 3.3.U5 Orientation of pairs of homologous chromosomes prior to separation is random.

Metaphase I
The bivalents line up at the equator.

Random orientation occurs - each

bivalent aligns independently and
hence the daughter nuclei get a
different mix of chromosomes.

This is a significant source of

genetic variation: there are 2n
possible orientations in metaphase
I and II. That is 223 in humans – or
8,388,068 different combinations in
gametes!
Edited from: http://www.slideshare.net/gurustip/meiosis-ahl courtesy of: http://www.flickr.com/carolinabio
10.1.U7 Independent assortment of genes is due to the random orientation of pairs of homologous
chromosomes in meiosis I.

Metaphase I
Allele have a 50 percent chance of moving to a
particular pole.
The direction in which one bivalent aligns does not
affect the alignment of other bivalents.
Therefore different allele combinations should always
be equally possible (if the gene loci are on different
chromosomes – this does not hold for linked genes)

Edited from: http://www.slideshare.net/gurustip/meiosis-ahl

3.3.S1 Drawing diagrams to show the stages of meiosis resulting in the formation of four haploid cells.

Anaphase I
Homologous pairs are separated and pulled to
opposing poles.

Spindle fibres contract.

This is the reduction division – the

bivalent is split and half the
chromosomes move towards each
pole.

Edited from: http://www.slideshare.net/gurustip/meiosis-ahl courtesy of: http://www.flickr.com/carolinabio

10.1.U5 Homologous chromosomes separate in meiosis I.

In anaphase I, the homologous pair is separated but

the sister chromatids remain attached.

This is the reduction division.

3.3.S1 Drawing diagrams to show the stages of meiosis resulting in the formation of four haploid cells. AND 3.3.U6
Separation of pairs of homologous chromosomes in the first division of meiosis halves the chromosome number.

Telophase I
The nuclei are now haploid (N) not diploid
(2N): they each contain one pair of sister
chromatids for each of the species’
chromosomes.

The cytoplasm begins to divide by

cytokinesis.

New nuclei form

Chromosomes decondense

n.b. If crossing-over and recombination

has occurred then the sister
chromatids will not be exact copies.

Edited from: http://www.slideshare.net/gurustip/meiosis-ahl courtesy of: http://www.flickr.com/carolinabio

3.3.S1 Drawing diagrams to show the stages of meiosis resulting in the formation of four haploid cells.

Prophase II

Nuclear membranes dissolves

Chromosomes, consisting of two sister
chromatids, condense

No crossing-over occurs.

centrioles move to opposite poles

(perpendicular to previous

Edited from: http://www.slideshare.net/gurustip/meiosis-ahl courtesy of: http://www.flickr.com/carolinabio

Review: 3.3.S1 Drawing diagrams to show the stages of meiosis resulting in the formation of four haploid cells.

Metaphase II
Pairs of sister chromatids align at the equator.

Spindle fibres form and attach at the

centromeres.

Edited from: http://www.slideshare.net/gurustip/meiosis-ahl courtesy of: http://www.flickr.com/carolinabio

10.1.U6 Sister chromatids separate in meiosis II.

Metaphase II
Pairs of sister chromatids align at the
equator. Spindle fibres form and
attach at the centromeres.

Random orientation again

contributes to variation in the
gametes, though not to such an
extent as in metaphase I.

This is because there is only a

difference between chromatids
where crossing-over has taken place.
Review: 3.3.S1 Drawing diagrams to show the stages of meiosis resulting in the formation of four haploid cells.

Anaphase II
The sister chromatids are separated. The
chromatids (now called chromosomes are
pulled to opposing poles.

Spindle fibres contract and the

centromeres are split.

Edited from: http://www.slideshare.net/gurustip/meiosis-ahl courtesy of: http://www.flickr.com/carolinabio

10.1.U6 Sister chromatids separate in meiosis II.

Telophase II
n.b. due to crossing over each of
the four new nuclei is likely to be
genetically different.

New haploid nuclei are formed.

Cytokinesis begins, splitting the

cells.

The end result of meiosis is four

haploid gamete cells.

Fertilisation of these haploid

gametes will produce a diploid
zygote.
Which phase of meiosis is shown? Why?
A. Interphase

B. Prophase I

C. Metaphase I

D. Metaphase II

Reason:
Which phase of meiosis is shown? Why?
A. Interphase

B. Prophase I

C. Metaphase I

D. Metaphase II

Reason:
• Homologous pairs are aligned (at equator), so must
be metaphase.
• Crossing-over has already taken place, so must be
after prophase I.
• Homologous pairs have not yet separated, so must
be still in meiosis I (metaphase I).
Which phase of meiosis is shown? Why?
A. Interphase

B. Prophase I

C. Metaphase I

D. Metaphase II

Reason:
Which phase of meiosis is shown? Why?
A. Interphase

B. Prophase I

C. Metaphase I

D. Metaphase II

Reason:
• Homologous pairs have associated.
• Crossing-over has taken place.
• Homologous pairs have not aligned at the equator.
Which phase of meiosis is shown? Why?
A. Interphase

B. Prophase I

C. Metaphase I

D. Metaphase II

Reason:

40
Which phase of meiosis is shown? Why?
A. Interphase

B. Prophase I

C. Metaphase I

D. Metaphase II

Reason:
• Homologous pairs have not yet associated.
• Replication has taken place.
• Crossing-over has not yet taken place.

41
Check your language. This image shows…

A. Four separate chromosomes.

B. A bivalent.

C. One pair of sister chromatids.

D. Non-disjunction.
Check your language. This image shows…

A. Four separate chromosomes.

B. A bivalent.

C. One pair of sister chromatids.

D. Non-disjunction.
Check your language. This image shows…

A. Two separate chromosomes.

B. A bivalent.

C. One pair of sister chromatids.

D. Crossing-over.
Check your language. This image shows…

A. Two separate chromosomes.

B. A bivalent.

C. One pair of sister chromatids.

D. Crossing-over.
Check your language. This image shows…

A. Two separate chromosomes.

B. A bivalent.

C. One pair of sister chromatids.

D. Homologous chromosomes.
Check your language. This image shows…

A. Two separate chromosomes.

B. A bivalent.

C. One pair of sister chromatids.

D. Homologous chromosomes.
Check your language. This image shows…

A. 8 separate chromosomes.

B. Two bivalents.

C. Two pairs of sister chromatids.

D. Two homologous chromosomes.

48
Check your language. This image shows…

A. 8 separate chromosomes.

B. Two bivalents.

C. Two pairs of sister chromatids.

D. Two homologous chromosomes.

Review: 3.3.U7 Crossing over and random orientation promotes genetic variation.

Prophase I Metaphase I Metaphase II

Crossing-over between non- Random orientation of the homologous chromosomes

sister chromatids results in means there are 2n possible orientations in metaphase I
recombination of alleles and II. That is 223 in humans – or 8,388,068 different
combinations in gametes!

Because both crossing-over and random orientation occur during meiosis the result is is
effectively infinite genetic variation in the haploid gamete.
n.b. for a new organism to arise sexually meiosis occurs in both parents further increasing
the genetic variation
Review: 3.3.U7 Crossing over and random orientation promotes genetic variation.

Prophase I Metaphase I Metaphase II

o r M eio sis II,

ro c ess, Meiosis I ive evidence
th e p n ? G
r t of riatio
Which pa e most genetic va
th
produces our answer.
rt y
to suppo

Crossing-over between non- Random orientation of the homologous chromosomes

sister chromatids results in means there are 2n possible orientations in metaphase I
recombination of alleles and II. That is 223 in humans – or 8,388,068 different
combinations in gametes!

Because both crossing-over and random orientation occur during meiosis the result is is
effectively infinite genetic variation in the haploid gamete.
10.1.U7 Independent assortment of genes is due to the random orientation of pairs of homologous
chromosomes in meiosis I.

Mendel’s Law of Independent Assortment

“The presence of an allele of one of the
A and B are different genes on
genes in a gamete has no influence over different chromosomes.
which allele of another gene is present.”
A is dominant over a.
B is dominant over b.

This only holds true for unlinked genes

(genes on different chromosomes).
10.1.U7 Independent assortment of genes is due to the random orientation of pairs of homologous
chromosomes in meiosis I.

Random Orientation vs Independent Assortment

“The presence of an allele of one of the
genes in a gamete has no influence over
which allele of another gene is present.”
Random Orientation refers to the
behaviour of homologous pairs of
chromosomes (metaphase I) or pairs of
sister chromatids (metaphase II) in
meiosis.
Independent assortment refers to the
behaviour of alleles of unlinked genes as
a result of gamete production (meiosis).

Due to random orientation of the chromosomes in

metaphase I, the alleles of these unlinked genes have
become independently assorted into the gametes.

Animation from Sumanas:

http://www.sumanasinc.com/webcontent/animations/content/independentassortment.html
10.1.U7 Independent assortment of genes is due to the random orientation of pairs of homologous
chromosomes in meiosis I.

Mendel and Meiosis “The presence of an allele of one of the genes in a gamete has no
influence over which allele of another gene is present.”

Mendel deduced that characteristics were determined by the interaction

between pairs of alleles long before the details of meiosis were known.

Where Mendel states that pairs of alleles of a gene separate independently

during gamete production, we can now attribute this to random orientation of
chromosomes during metaphase I.
Mendel made this deduction when working with
pea plants. He investigated two separate traits
(colour and shape) and performed many test
crosses, recording the ratios of phenotypes
produced in subsequent generations.

It was rather fortunate that these two traits

happened to be on separate chromosomes
(unlinked genes)! Remember back then he did not
know about the contents of the nucleus.
Chromosomes and DNA were yet to be discovered.

We will use his work as an example of dihybrid

crosses in the next section.

Animation from Sumanas: http://www.sumanasinc.com/webcontent/animations/content/independentassortment.html

Nature of Science: Making careful observations—careful observation and record keeping turned up anomalous data that
Mendel’s law of independent assortment could not account for. Thomas Hunt Morgan developed the notion of linked genes
to account for the anomalies. (1.8)

Morgan’s experiments (1909 - 1914) with fruit flies Thomas Hunt Morgan developed
produced results that could not be explained by the idea of sex-linked genes
Mendel’s work on heredity as it stood.

The ‘anomalous’ data was repeated and found

to be predictable. The experiments lead
Morgan and his colleagues to revise Mendelian https://www.dnalc.org/view/15005-Thomas-
Hunt-Morgan.html
heredity (1915) to include certain key tenets:
• Discrete pairs of factors are located on
chromosomes (later to be called genes)
• Certain characteristics are sex-linked
• Other characteristics are also sometimes
associated

https://geneticsandevolutionch10.files.wordpress.com/2015/01/kbtnk9dz-13672096431.jpg?w=672&h=327
Columbia University Fly Room http://www.nature.com/scitable/content/ne0000/ne0000/ne0000/ne0000/122977784/1_2.jpg
Bibliography / Acknowledgments

Bob Smullen