LES106 Biology: Form and Function

Semester 1, 2023/2024
Week 4

Sammy Chan
Module Instructor
 This lecture
• Eukaryotic cell division
• Mitosis – growth, repair, regeneration (last lecture)
• Meiosis – sexual life cycle and reproduction
 Learning Objectives
Sexual Reproduction Requires Meiosis
• Characterize the function of meiosis in sexual reproduction
• Distinguish between germ-line and somatic cells

Features of Meiosis
• Describe how homologous chromosomes pair during meiosis
• Explain why meiosis I is called the reductive division
 Bacterial Cell Division
• Bacteria divide by binary fission
– No sexual cycle
– Asexual reproduction: used by single-cell
organisms, usually prokaryotes
– Reproduction is clonal (all cells identical to
parent)
 Essence of sexual reproduction
• The genetic contribution of two cells (gametes)
• Forms the initial cell of an organism
• Plays a key role in generating genetic diversity

Haploid sperm Paternal

Fertilization homologue
Maternal
homologue

Diploid zygote

Haploid egg
Products of meiosis Initial cell Sexual life cycle
Haploid sperm
Paternal
homologue
Fertilization
Maternal
homologue
Haploid egg Diploid zygote

Haploid sperm Paternal

Fertilization homologue
Maternal
homologue

Diploid zygote

Haploid egg Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
 Sexual life cycle
• Meiosis and fertilization constitute a cycle of reproduction
– Diploid cells (2n)
• Somatic cells of adults have 2 sets of chromosomes
– Haploid cells (n)
• Gametes have only 1 set of chromosomes

Gametes

Haploid sperm
Paternal
homologue
Fertilization
Maternal
homologue
Haploid egg Diploid zygote
 Sexual life cycle
• Allows offspring to inherit genetic material from 2 parents
e.g. humans inherit 23 chromosomes from the mother (maternal homologue)
and 23 chromosomes from the father (paternal homologue)

Gametes

Haploid sperm
Paternal
homologue
Fertilization
Maternal
homologue
Haploid egg Diploid zygote
 Sexual life cycle
• Life cycles of sexually reproducing organisms involve the alternation of
haploid and diploid stages

• Some life cycles include longer diploid phases, some include longer
haploid phases

Gametes

Haploid sperm
Paternal
homologue
Fertilization
Maternal
homologue
Haploid egg Diploid zygote
 Gamete
Sexual life cycle (haploid) n

Gamete
• In most animals, diploid state (haploid) n
n
dominates 2n

– Zygote first undergoes mitosis

to produce diploid cells
(zygote develops into foetus Zygote
(diploid) 2n
→ child → adult)
MITOSIS
– Later in the life cycle, some of
these diploid cells undergo
Somatic
meiosis to produce haploid Germ-line cells
cells Germ-line
gametes (sperm & eggs) cells

MITOSIS
Adult male
(diploid) 2n Adult female
(diploid) 2n
 To go from haploid to diploid,
Sperm chromosome number must
(haploid) n be doubled!
Meiosis: - Through fertilization
Produce haploid
gametes from Egg Haploid Stage
diploid parent cells (haploid) n n = 23
n
2n

To go from diploid to Diploid Stage

haploid, chromosome n = 46
number must be halved!
- Through meiosis Zygote
(diploid) 2n
MITOSIS

Mitosis:
Somatic Growth and
Germ-line cells
cells Germ-line
development in the
cells diploid stages
(zygote, child, adult)
MITOSIS
Adult male
(diploid) 2n Adult female
(diploid) 2n
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 Features of Meiosis
• Meiosis includes two rounds of division (PMAT)
– Meiosis I and Meiosis II
– Each has prophase, metaphase, anaphase, and telophase stages
 Features of Meiosis
• Meiosis I: termed “reductive division”
– Separates the homologous pairs of chromosomes
– Results in daughter cells that contain one homologue from each
homologous pair (coming from each parent)

• No DNA replication between meiotic divisions:

– DNA replicates once, cells divide twice → create haploid cells

• Meiosis II: does not further reduce the number of chromosomes

– Separates the sister chromatids for each homologue
 Features of Meiosis

Meiosis I
– Separates the homologous pairs of chromosomes

Meiosis II
– Separates the sister chromatids of each homologue
 Interphase
Homologous pair of chromosomes
in diploid parent cell

Before Meiosis
Diploid: 2n
Chromosomes
replicate
S-phase of
Homologous pair of replicated chromosomes
Interphase
Sister
chromatids Diploid cell with
replicated
chromosomes

After Meiosis I Meiosis I

Meiosis I first
separates
Haploid cells
Homologous Pairs
with 1 Homologous

replicated
chromosomes of chromosomes
separate
chromosomes Haploid cells with
replicated chromosomes
Meiosis II Meiosis II then
2 Sister chromatids
separate
separates pairs of
Sister Chromatids
Haploid: n into individual
Haploid cells with unreplicated chromosomes
Chromosomes
 Learning Objectives
The Process of Meiosis
• Describe the key events that occur in Prophase I
• Describe arrangement of homologous pairs at metaphase I
• Differentiate between the events of anaphase I and anaphase II of meiosis
• Know how errors can occur in meiosis and the outcome of those errors
• Understand how meiosis results in genetic diversity
 The Process of Meiosis
• Meiotic cells have an interphase • Meiosis I
period that is similar to mitosis with – Prophase I
G1, S, and G2 phases – Metaphase I
– Anaphase I
• After interphase, germ-line cells – Telophase I
(cells that produce gametes) enter • Meiosis II
meiosis I – Prophase II
– Metaphase II
– Anaphase II
– Telophase II
 Meiosis I

Chromosome
replication

Meiosis I
• Chromosome replication prior to meiosis I
• Homologous pairs separate
• Genetic variation: crossing over in prophase I
 Prophase I
• Chromosomes become visible,
nuclear envelope breaks down,
spindle forms

• Each chromosome composed of

2 sister chromatids

• Homologous chromosomes
become closely paired
Synapsis (chromosome
pairing) Prophase I
• Homologous chromosomes
become closely associated
during prophase I
• Includes the formation of
synaptonemal complex Kinetochore

– Homologues Sister chromatids

Synaptonem
closely paired along a al
complex
lattice of proteins Homologues
between them (Non-sister
Chromatids)
Centromere

– Along the entire length

of the chromosomes Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

– Structure also
called tetrad or bivalents
 Prophase I
• Crossing over: exchange of DNA
between non-sister chromatids of
homologous pairs of chromosomes
→ Genetic variation

• Non-sister chromatids remain attached

at chiasmata
 Crossing Over
• Occurs between non-sister chromatids
Cohesin proteins Site of crossover

• Allows the maternal and paternal

homologues to exchange chromosomal chiasma

material

• Also called genetic recombination

chiasma

Prophase I Metaphase I

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 Crossing Over
• Alleles of genes that were formerly on
Cohesin proteins Site of crossover
separate homologues can now be found
on the same homologue
– Allele: version of a gene chiasma
E.g. Gene for eye color, alleles are brown or blue

• Chiasma – site of crossing over

– Plural: chiasmata chiasma

Prophase I Metaphase I

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 Crossing Over
• Each chromosome arm usually has one or
Cohesin proteins Site of crossover
a few crossovers per meiosis

• Humans typically have two or three chiasma

• Process increases genetic variation by

giving rise to new combinations of
genetic material within a species chiasma

Prophase I Metaphase I

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 Crossing Over
• The synaptonemal complex breaks down
Cohesin proteins Site of crossover
when crossing over is complete

• Chromosomes become less tightly chiasma

associated but remain attached by
chiasmata (until anaphase I)

chiasma

Prophase I Metaphase I

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 Crossing Over
• Thus, the 4 chromatids are held together
Cohesin proteins Site of crossover
in two ways:
1. The two sister chromatids of each
homologue (products of DNA chiasma
replication) are held together by
cohesin proteins, along the
chromosome arms (compare this
with mitosis)
chiasma
2. The two homologous chromosomes
are held together at chiasmata
Prophase I Metaphase I

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 Metaphase I
• Terminal chiasmata hold homologous pairs
together following crossing over

• Homologous pairs of chromosomes are

aligned at the metaphase plate side-by-side
• Aligned as paired homologues, not
individual chromosomes

• Orientation of each pair of homologues pairs

on the spindle is random
• Either member of the pair can be on the
left or right side of plate

• This sets up independent assortment

Homologues pairs of chromosomes align randomly at metaphase plate
• Many random combinations of the 3 pairs of homologous chromosomes

Example:
n=3
2n = 6

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 Genetic Variation in Meiosis:
1. Crossing over in Prophase 1

2. Independent Assortment in
Metaphase I

• Humans have 23 pairs; 223 (8,388,608) combinations are possible

from independent assortment alone
 Metaphase I Meiosis I
Metaphase I Crossovers and
sister chromatid
cohesion lock
• Microtubules from opposite poles attach homologues
to the kinetochores of each homologue together.
Microtubules
(not each sister chromatid) connect to the
kinetochores of
sister chromatids so
• i.e. microtubules from one pole attaches that homologues are
to one homologue, while microtubules pulled toward
Anaphase I opposite poles.
from the opposite pole attaches to the
other homologue of the same pair
 Anaphase I

• Microtubules of the spindle shorten

• Chiasmata break

• Homologous pairs of chromosomes are

separated from each other and move to
opposite poles
• Note: sister chromatids remain attached
to each other at their centromeres
 Anaphase I
Cohesin proteins Site of crossover

chiasma
chiasmata
break

chiasma

Prophase I Metaphase I Anaphase I

• Maternal and paternal homologues to exchange chromosomal material

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Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Meiosis I
Metaphase I Crossovers and
sister chromatid
cohesion lock
homologues
together.
Microtubules
connect to the
kinetochores of
sister chromatids so
that homologues are
pulled toward
Anaphase I opposite poles.

Microtubules pull
the homologous
chromosomes
apart, but sister
chromatids are
held together at
the centromere.
 Anaphase I

• Each pole now has a complete

haploid set of chromosomes
consisting of one member of each
homologous pair
 Telophase I
• Nuclear envelope re-forms around each
daughter nucleus

• Sister chromatids are no longer identical

because of crossing over (prophase I)

• Cytokinesis may or may not occur after

telophase I

• Meiosis II occurs after an interval of

variable length
 The Stages of Meiosis I Half the number of
chromosomes as in
parent cell

Chromosome
replication

• Prophase I: homologous chromosomes align, crossing over occurs at chiasmata

• Metaphase I: homologous pairs line up at metaphase plate
• Anaphase I: pairs of homologous chromosomes separate, move to opposite poles,
sister chromatids remain attached
• Telophase I (and cytokinesis): two haploid daughter cells, each containing one
homologue of each replicated chromosome (with sister chromatids)
 Meiosis II
• Resembles a mitotic division (PMAT)
– But with a haploid rather than diploid cell
 Prophase II
• Following a brief interphase, with no S
phase, meiosis II begins

• New spindle forms in each cell, nuclear

envelope breaks down
o In some species, nuclear envelope does
not reform in telophase I, this step is
obviated

• Chromosomes (each still composed of two

sister chromatids) move toward the
metaphase plate
 sister chromatids
Metaphase II

• Chromosomes align on metaphase plate

• Kinetochore microtubules from opposite poles

attach to kinetochores of sister chromatids, as
in mitosis
Recall from our last lecture
(mitosis)… Kinetochores

• Centromere – point of
Sister
Cohesin chromatids constriction
proteins

Centromere
• Kinetochore proteins
region of
chromosome
assemble. Attachment site for
microtubules
• Each sister chromatid has
a centromere
Kinetochore
Kinetochore
• Chromatids stay attached at
microtubules centromere by cohesin
Metaphase
chromosome

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 Anaphase II

• When microtubules shorten in anaphase II,

sister chromatids are separated and pulled
to opposite poles of the cells, as in mitosis

• Removal of cohesin proteins

 Telophase II

• Nuclear envelopes re-form around four

different clusters of chromosomes

• After cytokinesis, four haploid cells result

• No two cells are alike due to the

1. Random alignment of homologous pairs at
metaphase I (independent assortment)
2. Crossing over during prophase I
 Meiosis II

• Prophase II: nuclear envelopes dissolve and new spindle apparatus forms, chromosomes
(each still composed of two sister chromatids) move toward the metaphase plate
• Metaphase II: chromosomes align on metaphase plate
• Anaphase II: sister chromatids are separated from each other, individual chromosomes
move towards opposite poles
• Telophase II: nuclear envelope re-forms around 4 sets of daughter chromosomes;
cytokinesis follows
Meiosis Summary What we
end with:
What we Four haploid daughter cells
(each n)
started with:
One diploid parent cells
(each 2n)

Meiosis I Meiosis II

Homologous
chromosomes

Homologous chromosomes Sister chromatids

have separated have separated
 Final result of Meiosis
• Four cells containing haploid sets of chromosomes

• In animals, these cells develop directly into gametes

– In Spermtogenesis: 4 sperm
– In Oogenesis: 1 egg, 3 polar bodies

• In plants, fungi, and many protists, haploid cells then divide

mitotically
– Produce larger number of gametes
Chromosome
replication
S-Phase Overview of Meiosis I and II
Label
• Homologous pairs
• Sister chromatids
• Show crossing over

Meiosis I

Meiosis II
 Errors in Meiosis

• Nondisjunction
• Aneuploid gametes
• Most common cause of spontaneous abortion in humans
 Errors in Meiosis

• Nondisjunction:
failure of
chromosomes to
move to opposite
poles during either
meiotic division

http://bio1151.nicerweb.com/Locked/media/ch15/nondisjunction.html
 Errors in Meiosis

• Aneuploid gametes:
gametes with missing
or extra chromosomes

Down syndrome
• Having an extra chromosome: trisomy
Usually fatal, with some exceptions
 Aneuploidy

Turner’s syndrome Down syndrome

• Missing one chromosome: monosomy • Having an extra chromosome: trisomy
Embryos do not survive in most cases Usually fatal, with some exceptions

https://www.differencebetween.com/difference-between-monosomy-and-vs-trisomy/
 Learning Objectives
Meiosis Versus Mitosis
• Compare/contrast meiosis with mitosis

• Explain the importance of the suppression of replication between

meiotic divisions
 Meiosis vs. Mitosis
Meiosis is characterized by 4 features:

1. Homologous pairing and crossing over joins maternal and paternal

homologues during meiosis I

2. Sister chromatids remain connected at their centromeres throughout

meiosis I, and segregate together during anaphase I

3. Kinetochores of sister chromatids attach to the same pole in meiosis I and

opposite poles in meiosis II, mitosis

4. DNA replication is suppressed between meiosis I and meiosis II

1. Homologous pairing is specific to meiosis

Pairing of homologous chromosomes during propose I

– First deviation from mitosis
– Sets the stage for all subsequent differences
• How homologues find each other and become aligned: mystery of meiosis
• Some cytological evidence: telomers and other specific sites – necessary for pairing
• Exact process not clear

Sister chromatid cohesion

• Cohesion along chromosome arms
• Cohesin proteins here are meiosis-specific
1. Homologous pairing is specific to meiosis

Recombination
• Molecular details of crossing over are complex
• Process initiated by a double-stranded break (DSB) in one homologue
• Similar machinery in meiotic recombination and DSB repair
• Important to proper disjucntion
 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

MEIOSIS I
Prophase I Metaphase I Anaphase I Telophase I

Parent cell
(2n) Paternal
homologue
Chromosome
replication
Paired homologous
Homologous chromosomes pair; chromosomes Homologous chromosomes separate;
synapsis and crossing over occur. align on metaphase plate. sister chromatids remain together.

Chromosome MITOSIS
replication
Prophase Metaphase Anaphase Telophase
Homologous
chromosomes
Maternal
homologue

Two daughter cells

(each 2n)

Homologous chromosomes Individual homologues align

Sister chromatids separate, cytokinesis occurs, and two cells
do not pair. on metaphase plate.
result, each containing the original number of homologues.

• In Mitosis (bottom), chromosomes align individually in single file

• Result: each daughter cell gets one copy of every single chromosome (all 46 = diploid)

• In Meiosis (top): chromosomes align as homologous pairs

• Result: each daughter cell gets only one member of the pair (23 = haploid)
2. Sister chromatid cohesion is maintained through meiosis I but released
in meiosis II
• Meiosis I characterised by separation of homologues, not sister chromatids, during anaphase I
• Sister chromatids must co-segregate (move towards the same pole)
• Sister chromatids are held together by cohesin proteins along their arms
• Homologues are joined by chiasmata
• Sister chromatid cohesion around site of exchange then holds the homologues together
• Anaphase I: destruction of cohesin along the arms
→ loss of chromosome arms cohesion
→ allows the homologues to be pulled apart at anaphase I
Cohesin proteins Site of crossover

chiasma chiasmata
break

chiasma

Prophase I Metaphase I Anaphase I

3. Sister kinetochores are attached to the same pole during meiosis I

• Co-segregation requires that the kinetochores of sister chromatids are attached to the same pole
during meiosis I – monopolar attachment
- In contrast to both mitosis and meiosis II where sister kinetochores become attached to opposite
poles
• Due to structural differences? Meiotic kinetochores seem to protrude more, monopolar attachment
easier. Mitotic kinetochores seem to be more recessed

• Sister chromatid cohesion at centromere and monopolar attachment are required for segregation of
homologues
Meiosis I Mitosis
Metaphase I
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Meiosis I Mitosis
Metaphase I Crossovers and Metaphase Homologues do
sister chromatid not pair;
cohesion lock kinetochores of
homologues sister chromatids
together. remain separate;
Microtubules microtubules
connect to the attach to both
kinetochores of kinetochores on
sister chromatids so opposite sides of
that homologues are the centromere.
pulled toward
Anaphase I opposite poles. Anaphase

Microtubules pull Microtubules

the homologous pull sister
chromosomes chromatids
apart, but sister apart.
chromatids are
held together at
the centromere.
4. DNA replication is suppressed between meiotic divisions

• Mitotic division: after one division, new round of DNA replication must occur before
the next division
• Meiosis: chromosome number halved at meiosis I, but sister chromatids have not
yet separated. DNA replication between the two rounds of divisions must be
suppressed

• Cyclin B levels reduced

- Protein that checks DNA replication in cell cycle
- Results in failure to form initiation complexes for DNA replication to proceed
→ DNA replication supressed
 Meiosis produces cells that are not identical

Mitosis: daughter cells identical to parent cell

• Exact copying critical for growth, development, wound healing / regeneration

Meiosis
• Daughter cells (gametes) are rarely identical
• Due to random assortment of chromosomes at first meiotic division
– Gametes all carry entire haploid set of chromosomes, but within a set is a mixture of maternal
and paternal chromosomes
• Due to crossing over
– homologues have also exchanged genetic material
• Contributes to genetic diversity
 Advantages of sexual reproduction

• Asexual organisms – the only source of genetic variation is mutation

• Mutations that take place during the formation of germ cell lines are also a source
of variation in sexually reproducing organisms
• In contrast to mutation during asexual reproduction, these can be continually
reshuffled from one generation to the next
– when different parents combine their unique genomes
– the genes are mixed into different combinations by crossovers during prophase I and random
assortment at metaphase I
 Review: Meiosis I

Chromosome
replication

• Prophase I: homologous chromosomes align, crossing over occurs at chiasmata

• Metaphase I: homologous pairs line up at metaphase plate
• Anaphase I: pairs of homologous chromosomes separate, move to opposite poles,
sister chromatids remain attached
• Telophase I (and cytokinesis): two haploid daughter cells, each containing one
homologue of each replicated chromosome (with sister chromatids)
 Review: Meiosis II

• Prophase II: nuclear envelopes dissolve and new spindle apparatus forms, chromosomes
(each still composed of two sister chromatids) move toward the metaphase plate
• Metaphase II: chromosomes align on metaphase plate
• Anaphase II: sister chromatids are separated from each other, individual chromosomes
move towards opposite poles
• Telophase II: nuclear envelope re-forms around 4 sets of daughter chromosomes;
cytokinesis follows
 Question 1
Germ line cells undergo meiosis to form —
a. Gametes
b. Reproductive cells
c. Haploid cells
d. Sperm and egg
e. All of the above
 Question 2
Which of the following is an example of a somatic cell?
a. Skin cell
b. Liver cell
c. White blood cell
d. Cardiac muscle cell
e. All are somatic cells
 Question 3
Homologous chromosomes contain the same genes but are inherited from different
parents.
a. This is true
b. This is false
 Question 4
Genetic recombination occurs when —
a. Sister chromatids align gene for gene
b. Genes are exchanged between adjacent chromosomes
c. Homologous pairs line up across the metaphase plate
d. Chromosomes condense into heterochromatin
 Question 5
Homologous pairs of chromosomes align across the middle of the cell during
a. Metaphase I
b. Anaphase I
c. Metaphase II
d. Anaphase II
e. Telophase III
 Question 6
Reduction division occurs due to the lack of DNA replication between —
a. Prophase I and Metaphase I
b. Anaphase II and Telophase I
c. Telophase II and Prophase I
d. Telophase I and Prophase II
e. None of the above
 Question 7
A germ line cell with 36 total chromosomes produces __ gametes that will have ___
chromosomes each.
a. 2, 36
b. 2, 18
c. 4, 36
d. 4, 18
e. None of the above
 Question 8

If a mutation occurred in a somatic cell, would it be

passed on to the offspring?
a. It would be passed on
b. It would not be passed on
 Question 9
The four phases of meiosis II are essentially identical to the phases of mitosis.
a. This is true
b. This is false