CAMPBELL BIOLOGY IN FOCUS

URRY • CAIN • WASSERMAN • MINORSKY • REECE

10
Meiosis and
Sexual Life Cycles

Lecture Presentations by
Kathleen Fitzpatrick and
Nicole Tunbridge,
Simon Fraser University

© 2016 Pearson Education, Inc. SECOND EDITION

 Overview: Variations on a Theme

 Living organisms are distinguished by their ability to

reproduce their own kind
 Heredity is the transmission of traits from one
generation to the next
 Variation is demonstrated by the differences in
appearance that offspring show from parents and
siblings
 Genetics is the scientific study of heredity and
variation

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 Concept 10.1: Offspring acquire genes from parents
by inheriting chromosomes
 In a literal sense, children do not inherit particular
physical traits from their parents

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 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)

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  Most DNA is packaged into chromosomes
 For example, humans have 46 chromosomes in
their somatic cells, the cells of the body except for
gametes and their precursors
 Each gene has a specific position, or locus, on a
certain chromosome

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 Comparison of Asexual and Sexual Reproduction

 In asexual reproduction, a single individual passes

genes to its offspring without the fusion of gametes
 A clone is a group of genetically identical
individuals from the same parent
 In sexual reproduction, two parents give rise to
offspring that have unique combinations of genes
inherited from the two parents

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Figure 10.2

0.5 mm

Parent
Bud

(a) Hydra (b) Redwoods

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 Concept 10.2: 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

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 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
 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

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Figure 10.3

Technique Results
Pair of homologous
duplicated chromosomes

Centromere 5 m

Sister
chromatids
Metaphase
chromosome

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Figure 10.3-2
Results
Pair of homologous
duplicated chromosomes

Centromere 5 m

Sister
chromatids

Metaphase
chromosome

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Figure 10.3-3

5 m

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  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)
 Human males have one X and one Y chromosome
 The remaining 22 pairs of chromosomes are called
autosomes

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  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
 A diploid cell (2n) has two sets of chromosomes
 For humans, the diploid number is 46 (2n  46)

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  In a cell in which DNA synthesis has occurred, each
chromosome is replicated
 Each replicated chromosome consists of two
identical sister chromatids

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Figure 10.4

Key
Maternal set of
2n  6 chromosomes (n  3)
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)

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  A gamete (sperm or egg) contains a single set of
chromosomes and is haploid (n)
 For humans, the haploid number is 23 (n = 23)
 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

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 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

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  At sexual maturity, the ovaries and testes produce
haploid gametes
 Gametes are the only types of human cells
produced by meiosis rather than mitosis
 Meiosis results in one set of chromosomes in each
gamete
 Fertilization and meiosis alternate in sexual life
cycles to maintain chromosome number

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Figure 10.5
Key Haploid gametes (n  23)
Haploid (n)
Diploid (2n) Egg (n)

Sperm (n)

MEIOSIS FERTILIZATION

Testis
Ovary
Diploid
zygote
(2n  46)

Mitosis and
development
Multicellular diploid
adults (2n  46)
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 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

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  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

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Figure 10.6-1

Haploid (n)
Diploid (2n)

n Gametes n

MEIOSIS FERTILIZATION

Zygote
2n 2n
Diploid
multicellular Mitosis
organism

(a) Animals
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  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

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  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

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Figure 10.6-2

Haploid multi- Haploid (n)

cellular organism Diploid (2n)
(gametophyte)

Mitosis n Mitosis
n n
n n
Spores
Gametes

MEIOSIS FERTILIZATION

2n
2n
Diploid Zygote
multicellular
Mitosis
organism
(sporophyte)

(b) Plants and some algae

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  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

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Figure 10.6-3

Haploid unicellular or Haploid (n)

multicellular organism Diploid (2n)

Mitosis n Mitosis

n
n n
Gametes n

MEIOSIS FERTILIZATION

2n

Zygote
(c) Most fungi and some protists
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  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 contribute to genetic variation in
offspring

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 Concept 10.3: Meiosis reduces the number of
chromosome sets from diploid to haploid
 Like mitosis, meiosis is preceded by the duplication
of chromosomes
 Meiosis takes place in two sets of cell divisions,
called meiosis I and meiosis II
 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

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 The Stages of Meiosis

 For a single pair of homologous chromosomes in a

diploid cell, both members of the pair are duplicated
 The resulting sister chromatids are closely
associated all along their lengths
 Homologs may have different versions of genes,
each called an allele
 Homologs are not associated in any obvious way
except during meiosis

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Figure 10.7
Interphase
Pair of homologous
chromosomes in
diploid parent cell

Pair of duplicated Chromosomes

homologous duplicate
chromosomes

Sister
chromatids Diploid cell with
duplicated
chromosomes
Meiosis I

Homologous
chromosomes
separate

Haploid cells with

Meiosis II duplicated chromosomes
Sister chromatids
separate

Haploid cells with unduplicated chromosomes

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Figure 10.7-1

Interphase

Pair of homologous
chromosomes in
diploid parent cell

Pair of duplicated Chromosomes

homologous duplicate
chromosomes

Sister
chromatids Diploid cell with
duplicated
chromosomes

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Figure 10.7-2
Meiosis I

Homologous
chromosomes
separate

Haploid cells with

Meiosis II duplicated chromosomes

Sister chromatids
separate

Haploid cells with unduplicated chromosomes

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  Meiosis halves the total number of chromosomes
very specifically
 It reduces the number of sets from two to one, with
each daughter cell receiving one set of
chromosomes

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  In the first meiotic division, homologous pairs of
chromosomes pair and separate
 In the second meiotic division, sister chromatids of
each chromosome separate
 Four new haploid cells are produced as a result

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Figure 10.8

MEIOSIS I: MEIOSIS II:

Separates homologous chromosomes Separates sister chromatids
Telophase I Telophase II
Prophase I Metaphase I Anaphase I and Cytokinesis
Prophase II Metaphase II Anaphase II
and Cytokinesis

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Figure 10.8-1

MEIOSIS I: Separates homologous chromosomes

Prophase I Metaphase I Anaphase I Telophase I

and Cytokinesis

Centrosome Sister chromatids

Kinetochore remain attached
(with centriole pair) (at centromere)
Sister Chiasmata
chromatids Spindle Kinetochore
microtubules microtubules
Metaphase
plate

Cleavage
furrow
Fragments Homologous
Pair of of nuclear chromosomes
homologous envelope separate
chromosomes
Centromere

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Figure 10.8-1a

MEIOSIS I:
Separates homologous chromosomes

Prophase I Metaphase I

Centrosome Kinetochore
(with centriole pair) (at centromere)
Sister Chiasmata
chromatids Spindle Kinetochore
microtubules microtubules
Metaphase
plate

Fragments
Pair of of nuclear
homologous envelope
chromosomes
Centromere
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Figure 10.8-1b

MEIOSIS I:
Separates homologous chromosomes

Anaphase I Telophase I
and Cytokinesis

Sister chromatids
remain attached

Cleavage
furrow
Homologous
chromosomes
separate

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Figure 10.8-2

MEIOSIS II: Separates sister chromatids

Prophase II Metaphase II Anaphase II Telophase II

and Cytokinesis

Sister chromatids
separate Haploid daughter
cells forming

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 Prophase I
 Chromosomes condense progressively throughout
prophase I
 Homologous chromosomes pair up, aligned gene
by gene

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  In crossing over, nonsister chromatids exchange
DNA segments
 Each homologous pair has one or more X-shaped
regions called chiasmata
 Chiasmata exist at points where crossing over has
occurred.

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Metaphase I
 In metaphase I, homologous pairs 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 tetrad
 Microtubules from the other pole are attached to the
kinetochore of the other chromosome

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 Anaphase I
 In anaphase I, pairs of homologous chromosomes
separate
 One chromosome moves toward each pole, guided
by the spindle apparatus
 Sister chromatids remain attached at the
centromere and move as one unit toward the pole

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 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

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  In animal cells, a cleavage furrow forms; in plant
cells, a cell plate forms
 No chromosome duplication occurs between the
end of meiosis I and the beginning of meiosis II
because the chromosomes are already replicated

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  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

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 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

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 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

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 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

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 Telophase II and Cytokinesis
 Nuclei form, and the chromosomes begin
decondensing
 At the end of meiosis, there are four daughter cells,
each with a haploid set of unduplicated
chromosomes
 Each daughter cell is genetically distinct from the
others and from the parent cell

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 Crossing Over and Synapsis During Prophase I

 During prophase I, two members of a homologous

pair associate along their length, allele by allele
 A zipper-like structure called the synaptonemal
complex forms during this attachment (synapsis)
 DNA molecules of the maternal and paternal
chromatid are broken at matching points
 The DNA breaks are closed so that a paternal
chromatid is joined to a piece of maternal
chromatid, and vice versa

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Figure 10.9
Pair of homologous
DNA Centromere DNA chromosomes:
breaks breaks Paternal
Cohesins
sister
chromatids
Maternal
sister
Synaptonemal chromatids
complex forming

Synaptonemal
complex
Crossover
Crossover

Chiasmata

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Figure 10.9-1

Pair of homologous
DNA Centromere chromosomes:
DNA
breaks breaks Paternal
Cohesins sister
chromatids
Maternal
sister
chromatids
Synaptonemal
complex forming

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Figure 10.9-2

Synaptonemal
complex
Crossover
Crossover

Chiasmata

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 A Comparison of Mitosis and Meiosis

 Mitosis conserves the number of chromosome sets,

producing cells that are genetically identical to the
parent cell
 Meiosis reduces the number of chromosome sets
from two (diploid) to one (haploid), producing cells
that differ genetically from each other and from the
parent cell
 Meiosis includes two divisions after replication, each
with specific stages

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  Three events are unique to meiosis, and all three
occur in meiosis l
 Synapsis and crossing over in prophase I:
Homologous chromosomes physically connect and
exchange genetic information
 Alignment of homologous pairs at the metaphase
plate: Homologous pairs of chromosomes are
positioned there in metaphase I
 Separation of homologs during anaphase I

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  Sister chromatids stay together due to sister
chromatid cohesion
 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)

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Figure 10.10
MITOSIS MEIOSIS
Parent cell Chiasma MEIOSIS I
Prophase Prophase I
Pair of duplicated
Chromosome Chromosome homologs
Duplicated duplication duplication
chromosome 2n  6

Individual Pairs of
Metaphase chromosomes Metaphase I
chromosomes
line up. line up.

Anaphase Anaphase I
Sister chromatids Homologs
Telophase Telophase I
separate. separate.
Haploid
n3
Daughter
Sister cells of
chromatids meiosis I
MEIOSIS II
2n 2n separate.
Daughter cells n n n n
of mitosis Daughter cells of meiosis II
SUMMARY
Property Mitosis (diploid and haploid) Meiosis (diploid only)
DNA replication Occurs during interphase before mitosis begins Occurs during interphase before meiosis I begins
Number of divisions One, including prophase, prometaphase, Two, each including prophase, metaphase, anaphase, and telophase
metaphase, anaphase, and telophase
Synapsis of homologous Does not occur Occurs during prophase I along with crossing over between nonsister chromatids;
chromosomes resulting chiasmata hold pairs together due to sister chromatid cohesion

Number of daughter cells Two, each genetically identical to the parent Four, each haploid (n); genetically different from the parent
and genetic composition cell, with the same number of chromosomes cell and from each other

Role in the animal or Enables multicellular animal or plant Produces gametes (in animals) or spores (in the sporophyte plant);
plant body (gametophyte or sporophyte) to arise from a reduces number of chromosome sets by half and introduces
single cell; produces cells for growth, repair, genetic variability among the gametes or spores
and, in some species, asexual reproduction;
produces gametes in the gametophyte plant
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Figure 10.10-1

MITOSIS MEIOSIS
Parent cell Chiasma MEIOSIS I

Prophase Prophase I
Chromosome Chromosome
Duplicated Pair of duplicated
duplication 2n  6 duplication
chromosome homologs

Individual Pairs of
Metaphase chromosomes Metaphase I
chromosomes
line up. line up.

Anaphase Anaphase I
Sister chromatids Homologs
Telophase Telophase I
separate. separate.
Haploid
n3
Daughter
Sister cells of
chromatids meiosis I
MEIOSIS II
2n 2n separate.
Daughter cells n n n n
of mitosis Daughter cells of meiosis II

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Figure 10.10-1a

MITOSIS MEIOSIS
Parent cell Chiasma MEIOSIS I
Prophase Prophase I
Chromosome Chromosome Pair of
Duplicated duplicated
chromosome duplication 2n  6 duplication
homologs
Individual Pairs of
chromosomes chromosomes
Metaphase line up. Metaphase I
line up.

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Figure 10.10-1b

MITOSIS MEIOSIS
Anaphase Anaphase I
Telophase Homologs Telophase I
Sister chromatids
separate. separate.
Haploid
Sister Daughter n3
cells of
chromatids meiosis I
MEIOSIS II
2n 2n separate.
n n n n
Daughter cells
of mitosis Daughter cells of meiosis II

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Figure 10.10-2

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Figure 10.10-2a

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Figure 10.10-2b

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 Concept 10.4: 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

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 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
 Three mechanisms contribute to genetic variation
 Independent assortment of chromosomes
 Crossing over
 Random fertilization

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 Independent Assortment of Chromosomes

 Homologous pairs of chromosomes orient randomly

at metaphase I of meiosis
 In independent assortment, each pair of
chromosomes sorts maternal and paternal
homologs into daughter cells independently of the
other pairs

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  The number of combinations possible when
chromosomes assort independently into gametes is
2n, where n is the haploid number
 For humans (n = 23), there are more than 8 million
(223) possible combinations of chromosomes

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Figure 10.11-s1

Possibility 1 Possibility 2

Two equally probable

arrangements of
chromosomes at
metaphase I

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Figure 10.11-s2

Possibility 1 Possibility 2

Two equally probable

arrangements of
chromosomes at
metaphase I

Metaphase II

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Figure 10.11-s3

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

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 Crossing Over

 Crossing over produces recombinant

chromosomes, which combine DNA inherited
from each parent
 In meiosis in humans, on average, one to three
crossover events occur per chromosome pair

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  Crossing over contributes to genetic variation by
combining DNA, producing chromosomes with
new combinations of maternal and paternal alleles

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 Animation: Genetic Variation

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Figure 10.12-s1
Nonsister chromatids
Prophase I held together
of meiosis during synapsis
Pair of
homologs

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Figure 10.12-s2
Nonsister chromatids
Prophase I held together
of meiosis during synapsis
Pair of
homologs Synapsis and
Chiasma crossing over
site

Centromere

TEM

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Figure 10.12-s3
Nonsister chromatids
Prophase I held together
of meiosis during synapsis
Pair of
homologs Synapsis and
Chiasma crossing over
site

Breakdown of
Centromere proteins holding sister
chromatid arms together.
TEM
Anaphase I

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Figure 10.12-s4
Nonsister chromatids
Prophase I held together
of meiosis during synapsis
Pair of
homologs Synapsis and
Chiasma crossing over
site

Breakdown of
Centromere proteins holding sister
chromatid arms together.
TEM
Anaphase I

Anaphase II

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Figure 10.12-s5
Nonsister chromatids
Prophase I held together
of meiosis during synapsis
Pair of
homologs Synapsis and
Chiasma crossing over
site

Breakdown of
Centromere proteins holding sister
chromatid arms together.
TEM
Anaphase I

Anaphase II

Daughter
cells
Recombinant chromosomes
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Figure 10.12-1

Chiasma
site

Centromere

TEM

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 Random Fertilization

 Random fertilization adds to genetic variation

because any sperm can fuse with any ovum
(unfertilized egg)
 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

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  Crossing over adds even more variation
 Each zygote has a unique genetic identity

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 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

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  Asexual reproduction is less expensive than sexual
reproduction
 Nonetheless, sexual reproduction is nearly
universal among animals
 Overall, genetic variation is evolutionarily
advantageous

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Figure 10.6

Haploid (n) Haploid multi- Haploid unicellular or

Diploid (2n) cellular organism multicellular organism
(gametophyte)
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

Zygote MEIOSIS FERTILIZATION

2n 2n
2n
2n
Diploid Diploid Zygote 2n
multicellular Mitosis multicellular Mitosis
organism organism Zygote
(sporophyte)

(a) Animals (b) Plants and some algae (c) Most fungi and some protists

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Figure 10.UN01-1

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Figure 10.UN01-2

Budding yeast

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Figure 10.UN02

Prophase I: Each pair of homologous chromosomes

undergoes synapsis and crossing over between
nonsister chromatids with the subsequent appearance
of chiasmata.

Metaphase I: Chromosomes line up as homologous

pairs on the metaphase plate.

Anaphase I: Homologs separate from each other;

sister chromatids remain joined at the centromere.

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Figure 10.UN03

F
H

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Figure 10.UN04

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