Mitosis and Meiosis

A K A Mandal

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• When cells divide, the chromosomes must also make copies of
themselves (replicate) to maintain the appropriate
chromosome number in the descendant cells
• In eukaryotes, the chromosomes replicate in two main types of
nuclear divisions, called mitosis and meiosis
• Even though these two types of divisions are quite different
and have different functions, some of the molecular features
are held in common
• Three common molecular processes to focus on are DNA
replication, adhesion of replicated chromosomes, and
orderly movement of chromosomes into descendant cells

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 Mitosis
• Mitosis is the nuclear division associated with the asexual
division of cells
• In multicellular organisms mitosis takes place during the
division of somatic cells, the cells of the body
• In single-celled eukaryotes such as yeasts, mitosis occurs in
the cell divisions that cause population growth. Since asexual
cell division is aimed at straightforward reproduction of cell
type, it is necessary for the set of chromosomes to be
maintained constant down through the cell generations, and
this is what mitosis achieves

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• The stages of the cell division cycle are similar in most
organisms
• The two basic parts of the cycle are interphase (comprising
gap 1, synthesis, and gap 2) and mitosis
• An event essential for the propagation of
genotype takes place in the S phase (synthesis
phase) because it is here that the actual
replication of the DNA of each chromosome
occurs. As a result of DNA replication, each
chromosome becomes two side-by-side units
called sister chromatids. The sister chromatids stay attached
through the action of specific adherence proteins.
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Stages of the cell cycle

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6
• Interphase is the "resting" or non-mitotic
portion of the cell cycle. 
• It is comprised of G1, S, and G2 stages of the
cell cycle. DNA is replicated during the S phase
of Interphase

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• 1. Prophase:
• The pairs of sister chromatids, which cannot be seen during
interphase, become visible. The chromosomes contract into a
shorter, thicker shape that is more easily moved around
• The chromosomes condense and become visible
• The centrioles form and move toward opposite ends of the
cell ("the poles")
• The nuclear membrane dissolves
• The mitotic spindle forms (from the centrioles in animal cells)
• Spindle fibers from each centriole attach to each sister
chromatid at the kinetochore

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• 2. Metaphase: The sister chromatid pairs come to lie in the
equatorial plane of the cell
• The Centrioles complete their migration to the poles
• The chromosomes line up in the middle of the cell ("the
equator")

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• 3. Anaphase: The sister chromatids are pulled to opposite
ends of the cell by microtubules that attach to the centromeres.
The microtubules are part of the nuclear spindle, a set of
parallel fibers running from one pole of the cell to the other
Nuclear spindle fibers provide the motive force that pulls apart
the chromosomes or chromatids in mitosis and meiosis
• In nuclear division, spindle fibers form parallel to the cell axis,
connected to one of the cell poles. These spindle fibers are
polymers of a protein called tubulin. Each centromere acts as a
site to which a multiprotein complex called the kinetochore
binds

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• The kinetochore acts as the site for attachment to spindle
fiber microtubules. From one to many microtubules from one
pole attach to one kinetochore, and a similar number from the
opposite pole attach to the kinetochore on the homologous
chromatid. Although the microtubules of the spindle appear
like ropes, their action is not ropelike. Instead the tubulin
depolymerizes at the kinetochores, shortening the microtubule
and thereby pulling the sister chromatids apart

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• Later, the chromatids are further separated by
the action of molecular motor proteins acting
on another set of microtubules not connected
to the kinetochore but running from pole to
pole. The spindle apparatus and the complex
of kinetochores and centromeres are what
determine the fidelity of nuclear division

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• 4. Telophase: Chromatids have arrived at the
poles and the pulling-apart process is
complete. A nuclear membrane reforms
around each nucleus, and the cell divides into
two daughter cells. Each daughter cell inherits
one of each pair of sister chromatids, which
now become chromosomes in their own right.

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• Thus overall, the main events of mitosis are
replication and sister chromatid adhesion,
followed by segregation of the sister
chromatids into each daughter cell. In a
diploid cell, for any chromosomal type the
number of copies goes from 2 4 2.

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 Cytokinesis Divides the Cytoplasm
In animal cells, cytokinesis occurs by a process known as cleavage
• First, a cleavage furrow appears
– cleavage furrow = shallow groove near the location of the old metaphase plate
• A contractile ring of actin microfilaments in association with myosin, a
protein
• The contraction of a the dividing cell's ring of microfilaments is like the
pulling of drawstrings
– The cell is pinched in two

Cytokinesis in plant cells is different because plant cells have cell walls.
• There is no cleavage furrow
• During telophase, vesicles from the Golgi apparatus move along
microtubules to the middle of the cell (where the cell plate was) and
coalesce, producing the cell plate
– Cell-wall construction materials are carried in the vesicles and are continually deposited
until a complete cell wall forms between the two daughter cells
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 Meiosis
• Meiosis is the general name given to two successive nuclear
divisions called meiosis I and meiosis II. Meiosis takes place in
special diploid cells called meiocytes
• Because of the two successive divisions, each meiocyte cell
gives rise to four cells, 1 cell : 2 cells : 4 cells. The four cells are
called products of meiosis
• In animals and plants, the products of meiosis become the
haploid gametes
• In humans and other animals, meiosis takes place in the
gonads, and the products of meiosis are the gametes—sperm
(more properly, spermatozoa) and eggs (ova). In flowering
plants, meiosis takes place in the anthers and ovaries, and
the products of meiosis are meiospores, which eventually
give rise to gametes. 21
• Before meiosis, an S phase duplicates each
chromosome’s DNA to form sister chromatids,
just as in mitosis

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• Meiosis then proceeds through the following stages:
• 1. Prophase I: As in mitosis, the sister chromatids become
visible, closely adhered side by side. However, in contrast with
mitosis, the sister chromatids (although fully replicated at the
DNA level) show an apparently undivided centromere. The
sister chromatid pairs at this stage are called dyads, from the
Greek word for “two.”

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• 2. Metaphase I: The homologous dyads now pair to form
structures called bivalents. Thus, any one bivalent contains a
total of four chromatids, sometimes referred to as a tetrad
(Greek; four). This stage represents the most obvious
difference from mitosis.
• Pairing of the dyads to form a bivalent is accomplished by
molecular assemblages called synaptonemal complexes along
the middle of the tetrads

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• Nonsister chromatids of the tetrad engage in a breakage-and-
reunion process called crossing over. The crossover is visible
as two chromatids crossing each other to form a structure
called a chiasma (plural, chiasmata). For those species that
have been carefully studied, a minimum of one crossover per
tetrad is known to be a necessary prelude to subsequent
orderly chromosome separation.

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 Crossing Over

• Genetic material from

the homologous
chromosomes is randomly
swapped
• This creates four unique
chromatids
• Since each chromatid is
unique, the overall genetic
diversity of the gametes is
greatly increased

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• 3. Anaphase I: Each of the two pairs of sister
chromatids (dyads) is pulled to a different
pole.

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• 4. Telophase I: A nucleus forms at each pole.

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• 5. Prophase II: The dyads reappear
Centrioles form and move toward the poles.
The nuclear membrane dissolves

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• 6. Metaphase II: The dyads move to the
equatorial plane.
Microtubules grow from the centrioles and
attach to the centromeres

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• 7. Anaphase II: Each of the sister chromatids
of a dyad is pulled into a different daughter
nucleus as the cells divide for a second time.

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8. Telophase II:
The chromosomes may decondense (depends on
species)
Cytokinesis reaches completion, creating four
haploid daughter cells

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A Comparison between Mitosis and Meiosis

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• We see therefore that the fundamental events of
meiosis are DNA replication and sister chromatid
adhesion, followed by homologous pairing,
segregation, and then another segregation.
Hence, within a single cell, the number of copies
of a chromosome of the same type goes from 2 :
4 : 2 : 1, and each product of meiosis must
therefore contain one chromosome of each type,
half the number of the original meiocyte.

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• Even though early investigators did not know about DNA or
that it is replicated during interphase, it was still evident from
observing mitosis under the microscope that mitosis is the
way in which the chromosome number is maintained during
cell division. But the sex cycle still presented a puzzle. Two
gametes join in the fertilization event. The early investigators
knew that in this process two nuclei fuse but that the
chromosome number of the fusion product nevertheless is
the standard for that species. What prevents the doubling of
the chromosome number at each generation? This puzzle was
resolved by the prediction of a special kind of nuclear division
that halved the chromosome number. This special division,
which was eventually discovered in the gamete-producing
tissues of plants and animals, is called meiosis.

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 Chromosome behavior and
inheritance patterns in eukaryotes
The basic life cycles
• Eukaryotes have three basic types of life
cycles:
– Diplod
– Haploid
– The alternation of diploid and haploid stages in
the life cycle of plants

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• Diploids: organisms that are in the diploid state for most of
their life cycle; that is, for most of their life cycle they consist
of cells having two sets of homologous chromosomes.
Animals are examples of organisms with this type of life cycle;
in most species diploid cells arise from a fertilized egg.
Meiosis takes place in special diploid meiocytes set aside for
this purpose in the gonads (testes and ovaries) and results in
haploid gametes. These are the sperm and eggs that unite
when the egg is fertilized, producing the zygote. The zygote
subsequently goes through repeated mitotic division to
produce the multicellular state

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The diploid life cycle

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• Haploids: organisms that are in the haploid state for most
of their life cycle. Common examples are molds and
yeasts, both fungi. An organism arises as a haploid spore,
which then through mitotic divisions produces a branching
network of end-to-end haploid cells (as in molds) or a
population of identical cells (as in yeasts). How can meiosis
take place in a haploid organism? After all, meiosis
requires the pairing of two homologous chromosome sets.
The answer is that two haploid cells of two parental strains
fuse to form a temporary diploid meiocyte. Meiosis takes
place in the meiocyte, forming haploid spores.

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The haploid life cycle

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• Organisms with alternating haploid-diploid generations:
organisms that are haploid for part of the life cycle and diploid
for part of the life cycle. Both haploid and diploid parts grow
by mitosis, but meiosis occurs only in the diploid stage. Plants
show such alternation of haploid and diploid generations:
the organism during the haploid stage of the cycle is called
the gametophyte and during the diploid part the sporophyte.
Plants such as ferns and mosses have separate free-living
haploid and diploid stages. Flowering plants are
predominantly diploid but do have a small haploid
gametophytic stage parasitic on the diploid within the flower .
However, for most genetic purposes plants can be treated
simply as showing a diploid cycle.
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The alternation of diploid and
haploid stages in the life cycle of
plants

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 Meiosis Creates Genetic Diversity
• Sexually reproducing organisms inherit two full sets of
chromosomes, one from each parent. Each set contains
autosomes, which are common to all members of the species,
and sex chromosomes, which are differently distributed
according to the sex of the individual
• Therefore, each diploid nucleus contains two closely similar
versions of each autosome, plus a set of sex chromosomes
appropriate to the sex of the individual. The two copies of
each autosome, one from the mother and one from the
father, are called homologous chromosomes, or homologs,
and in most cells they maintain a separate existence as
independent chromosomes

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 Meiosis Creates Genetic Diversity

• During meiosis, however, each chromosome must

communicate with its unique homologous partner by
physically pairing and undergoing genetic recombination. This
communication is essential to enable the homologs to
segregate accurately into different daughter cells during
meiosis.

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• A crucial feature of meiosis is that it generates haploid cells
that are genetically different from one another and from the
two haploid cells that formed the organism in the first place.
• The genetic differences arise by two mechanisms:
– First, an individual gamete contains either the maternal or paternal
version of each chromosome; because the choice of maternal or
paternal occurs independently and randomly for each pair of
homologs, the original maternal and paternal chromosomes are
reshuffled into novel combinations in the haploid cells.
– Second, although the maternal and paternal versions of each
chromosome have similar DNA sequences, they are not identical, and
they undergo genetic recombination during meiosis—a process called
crossing-over to produce novel hybrid versions of each chromosome;
thus, each chromosome in a gamete contains a unique mixture of
genetic information from both parents

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 Duplicated Homologs (and Sex Chromosomes) Pair
During Early Prophase I

• During mitosis in most organisms, homologous

chromosomes behave independently of each
other. During meiosis I, however, it is crucial that
homologs recognize each other and associate
physically in order for the maternal and paternal
homologs to undergo genetic recombination
and to segregate to different daughter cells at
anaphase I. Special mechanisms mediate these
intimate interactions between homologs

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• The progressive juxtaposition of homologs occurs during a
very prolonged meiotic prophase (prophase I), which can take
hours in yeasts, days in mice, and weeks in higher plants.
• Like their mitotic counterparts, duplicated meiotic prophase
chromosomes initially appear as long threadlike structures, in
which the sister chromatids are so tightly glued together that
they appear as one. It is during early prophase I that the
homologs begin to associate along their length in a process
called pairing, which, in some organisms at least, occurs
initially through interactions between complementary DNA
sequences (called pairing sites) in the two homologs; in most
organisms, stable pairing requires genetic recombination
between the homologs
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• As prophase I progresses, the homologs
become more closely juxtaposed, forming a
four-chromatid structure called a bivalent
• Genetic recombination begins during pairing
in early prophase I, with the production of
programmed doublestrand breaks in
chromatid DNA; some of these
recombination events will later resolve into
crossovers, where a fragment of a maternal
chromatid is exchanged for a corresponding
fragment of a homologous paternal
chromatid.

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Homolog alignment and crossing-over.
• (A) The structure formed by two closely
aligned duplicated homologs is called a
bivalent. As in mitosis, the sister
chromatids in each homolog are tightly
connected along their entire lengths, as
well as at their centromeres. At this stage,
the homologs are usually joined together
by a protein complex called the
synaptonemal complex
• (B) A later-stage bivalent in which a single
crossover event has occurred between
non-sister chromatids. It is only when the
synaptonemal complex disassembles and
the paired homologs separate a little at
the end of prophase I, as shown, that the
crossover is seen microscopically as a thin
connection between the homologs called
a chiasma.
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• The pairing of homologs requires chromosome movements,
but it is not known what drives these movements
• The replicated chromosomes undergo major rearrangements
within the nucleus during prophase I. The ends of the
chromosomes (the telomeres) are tightly bound to the inner
surface of the nuclear envelope. They are initially distributed
diffusely there, but they then cluster transiently at one spot
on the envelope and, later still, disperse again.

Rearrangements of
telomeres during prophase in
developing bovine eggs. The
nucleus is
stained blue and the
telomeres are
stained red.
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• Neither the mechanism nor the roles of these rearrangements
are known, although they are thought to make prophase I
faster and more efficient. One possibility is that they help
prevent chromosome entanglements during prophase I.
• In fission yeast, telomere clustering is required for homolog
pairing and crossingover, but in some organisms it occurs after
pairing is well underway.

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 Pairing of Sex Chromosomes?
• We have described the pairing of homologous autosomes
during prophase I, but what happens to the sex
chromosomes?
• This varies between different organisms. Female mammals
have two X chromosomes, which pair and segregate like other
homologs. But the males have one X and one Y chromosome.

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• Although these chromosomes are not homologous, they too
must pair and undergo crossing-over during prophase if they
are to segregate normally at anaphase I. Pairing, crossing-
over, and segregation are possible because of a small region
of homology between the X and the Y at one or both ends of
these chromosomes
• The two chromosomes pair and crossover in this region
during prophase I, ensuring that each sperm receives either
one Y or one X chromosome and not both or neither
• Thus, only two types of sperm are normally produced: those
containing one Y chromosome, which will give rise to male
embryos, and those containing one X chromosome, which will
give rise to female embryos
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 Homolog Pairing Culminates in the Formation of a
Synaptonemal Complex

• The paired homologs are brought into closer juxtaposition,

with their structural axes (axial cores) about 400 nm apart, by
a mechanism that depends in most species on the
programmed double-strand DNA breaks that occur in sister
chromatids.

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 Homolog Pairing Culminates in the Formation of a
Synaptonemal Complex
• What pulls the axes together? One possibility is that the large
protein machine, called a recombination complex, which
assembles on a double-strand break in a chromatid, binds the
matching DNA sequence in the nearby homolog and helps
reel in this partner. This so-called
presynaptic alignment of the
homologs is followed by synapsis,
in which the axial core of a homolog
becomes tightly linked to the axial
core of its partner by a closely packed array of transverse
filaments to create a synaptonemal complex, which bridges
the gap, now only 100 nm, between the homologs .
Simplified
schematic
drawing of a
synaptonemal
complex.

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• The morphological changes that occur during the pairing of
meiotic chromosomes are the basis for dividing prophase I
into five sequential stages—leptotene, zygotene, pachytene,
diplotene, and diakinesis. Prophase I starts with leptotene,
when homologs condense and pair, and genetic
recombination begins. At zygotene, the synaptonemal
complex begins to assemble in local regions along the
homologs; assembly initiates at sites where the homologs are
closely associated and recombination events are occurring.

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Homolog synapsis and desynapsis during the different stages
of prophase I. (A) A single bivalent is shown schematically. At
leptotene, the two sister chromatids coalesce, and their
chromatid loops extend out together from a common axial
core. The synaptonemal complex begins to assemble locally in
early zygotene. Assembly continues through zygotene and is
complete in pachytene. The complex disassembles in
diplotene. 59
• At pachytene, the assembly process is complete, and the
homologs are synapsed along their entire lengths. The
pachytene stage can persist for days or longer, until
desynapsis begins at diplotene with the disassembly of the
synaptonemal complexes and the concomitant condensation
and shortening of the chromosomes.
• It is only at this stage, after the complexes have disassembled,
that the individual crossover events
between nonsister chromatids can be seen
as interhomolog connections called
chiasmata (singular chiasma), which now
play a crucial part in holding the compact
homologs together. The homologs are now ready to begin the
process of segregation. Prophase I ends with diakinesis—the
stage of transition to metaphase I. 60
A bivalent with three
chiasmata resulting
from three crossover
events. (A) Light
micrograph of a
grasshopper bivalent.
(B) Drawing showing
the arrangement of the
crossovers in (A).

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• The proteins that form the transverse filaments that bridge
between the axial cores of the homologs have been identified
in several species, including yeasts, worms, flies, and
mammals. They form homodimers that interact with each
other across the 100 nm gap between the homologs. In most
eucaryotes, these proteins are important for crossing-over,
as mutants that lack them fail to form crossovers. The cohesin
complexes that assemble on the DNA during S phase and bind
the sister chromatids together during meiosis are major
components of the axial core of each homolog. Some of the
cohesin subunits that operate in meiosis are the same as
those that function in mitosis, whereas others are specific for
meiosis. Both the crossovers and the cohesin complexes play
crucial parts in segregating the homologs during meiotic
division I 62
A molecular model of how
transverse filaments may be
formed by a single type of
protein

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 Homolog Segregation Depends on Meiosis-Specific,
Kinetochore-Associated Proteins
• One fundamental difference between meiosis I and mitosis
(and meiosis II) is that in meiosis I homologs rather than
sister chromatids separate and then segregate into the two
daughter cells. This difference depends on three features of
meiosis I that distinguish it from mitosis
• First, the kinetochores (protein complexes associated with the
centromeres) on the two sister chromatids of a homolog
attach to microtubules emanating from the same pole of the
meiosis I spindle and thus segregate together into the same
daughter cell at anaphase I; this contrasts with mitosis (and
meiosis II), in which the kinetochores on the two sister
chromatids of a chromosome attach to opposite poles of the
spindle and therefore segregate into different daughter cells
at anaphase. I. 64
 Homolog Segregation Depends on Meiosis-Specific,
Kinetochore-Associated Proteins

• Second, a strong physical linkage is maintained between the

homologs that resists the pulling forces of the meiosis I
spindle until the bivalents are aligned at the equator of the
spindle and the homologs separate at anaphase I. The
chiasmata formed between nonsister chromatids and the
cohesion between sister-chromatid arms cooperate in holding
the homologs together
• Third, the arms of the sister chromatids separate at anaphase
I, resolving the chiasmata and allowing the homologs to
separate, but the sisters stay glued together in the region of
their centromeres until anaphase II and therefore do not
separate in anaphase I.
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• Unlike meiosis I, meiosis II occurs rapidly and closely
resembles a mitotic cell division, although it occurs
without DNA replication. Prophase II is brief: the
nuclear envelope breaks down as the new spindle
forms, after which metaphase II, anaphase II, and
telophase II usually follow in quick succession. After
nuclear envelopes have formed around the four
haploid nuclei produced at telophase II, cytokinesis
occurs, and meiosis is complete.

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Comparison of chromosome behavior in
meiosis I, meiosis II, and mitosis.
Chromosomes behave similarly in mitosis and
meiosis II, but they behave very differently in
meiosis I. (A) In meiosis I, the two sister
kinetochores are located side-by-side on each
homolog at the sister centromeres and attach
to microtubules emanating from the same
spindle pole. The proteolytic destruction of the
cohesin complexes along the sister chromatid
arms unglues the arms and resolves the
crossovers, allowing the duplicated homologs to
separate at anaphase I, while the residual
cohesin complexes at the centromeres keep the
sisters together. The proteolytic destruction of
the residual cohesin complexes at the
centromeres allows the sister chromatids to
separate at anaphase II. (B) In mitosis, by
contrast, the two sister kinetochores attach to
microtubules emanating from different spindle
poles, and the two sister chromatids come
apart at the start of anaphase and segregate
into separate daughter cells 67
 Meiosis Frequently Goes Wrong
• The sorting of chromosomes that takes place during meiosis is
a remarkable feat of intracellular bookkeeping. In humans,
each meiosis requires that the starting cell keep track of 92
chromatids (46 chromosomes, each of which has duplicated),
distributing one complete set of each type of chromosome to
each of the four haploid progeny cells. Not surprisingly,
mistakes can occur in allocating the chromosomes during this
elaborate process. Mistakes are especially common in human
female meiosis, which arrests for years after diplotene:
meiosis I is completed only at ovulation, and meiosis II only
after the egg is fertilized. Indeed, such chromosome
segregation errors during egg development are the
commonest cause of both spontaneous abortion (miscarriage)
and mental retardation in humans. 68
• When homologs fail to separate properly—a phenomenon
called nondisjunction—the result is that some of the haploid
gametes produced lack a particular chromosome, while
others have more than one copy of it. (Cells with an abnormal
number of chromosomes are said to be aneuploid, whereas
those with the correct number are said to be euploid.) Upon
fertilization, aneuploid gametes form abnormal embryos,
most of which die. Some survive, however. Down syndrome in
humans, for example, which is the leading single cause of
mental retardation, is caused by an extra copy of
chromosome 21, usually resulting from nondisjunction during
meiosis I in the female ovary. Segregation errors during
meiosis I increase greatly with advancing maternal age.
69
• Despite its fallibility, almost all eucaryotes use
meiosis, intermittently at least, to shuffle their
genetic information before passing it on to the
next generation.
• Crossing-over makes a major contribution to
this genetic shuffling process.

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Meiosis – division error

Chromosome pair
Meiosis error - fertilization

Should the gamete with the

chromosome pair be fertilized then
the offspring will not be ‘normal’.

In humans this often occurs with the

21st pair – producing a child with
Downs Syndrome
21 trisomy – Downs Syndrome
 Crossing-Over Is Highly Regulated
• Crossing-over has two distinct functions in meiosis: it helps
hold homologs together so that they are properly segregated
to the two daughter cells produced by meiosis I, and it
contributes to the genetic diversification of the gametes that
are eventually produced. As might be expected, therefore,
crossing-over is highly regulated: the number and location of
double-strand breaks along each chromosome is controlled,
as is the likelihood that a break will be converted into a
crossover. Although the double-strand breaks that occur in
meiosis I can be located almost anywhere along the
chromosome, they are not distributed uniformly: they cluster
at “hot spots”, where the chromatin is accessible, and occur
only rarely in “cold spots”, such as the heterochromatin
regions around centromeres and telomeres. 74
• At least two kinds of regulation influence the location and
number of crossovers that form, neither of which is well
understood. Both operate before the synaptonemal complex
assembles. One ensures that at least one crossover forms
between the members of each homolog pair, as is necessary
for normal homolog segregation in meiosis I. In the other,
called crossover interference, the presence of one crossover
event inhibits another from forming close by, perhaps by
locally depleting proteins required for converting a double-
strand DNA break into a stable crossover.

75
 Meiosis Is Regulated Differently in Male and
Female Mammals

• The basic mechanisms of meiosis have been

conserved in evolution in all sexually reproducing
eucaryotes. In all of these organisms, for example,
most of meiosis is spent in prophase I, although the
details of the timing of different stages vary among
organisms. There are, however, some remarkable
differences in the regulation of meiosis in different
species and in different sexes of the same species.
The difference between the two sexes is very
striking in mammals
76
• In mammalian females, egg precursor cells (oocytes) begin
meiosis in the fetal ovary but arrest after diplotene, after the
synaptonemal complex has disassembled in meiosis I. They
complete meiosis I only after the female has become sexually
mature and the oocyte is released from the ovary during
ovulation; moreover, the released oocyte completes meiosis II
only if it is fertilized. Thus, there are special stop and start
mechanisms during meiosis in female mammals.
• In humans, some oocytes remain arrested in meiosis I for 40
years or more, which is presumably at least part of the reason
why nondisjunction increases dramatically in older women. In
mammalian males, by contrast, meiosis only begins in sperm
precursor cells (spermatocytes) in the testes at puberty and
then goes on continuously, without the stop and start
mechanisms that operate in female meiosis. It takes about 24
days for a human spermatocyte to complete meiosis. 77
• There is also a big difference in the error rates
of meiosis in mammalian females and males,
and this is especially striking in humans
• About 20% of human eggs are aneuploid,
compared with 3–4% of human sperm, and,
largely as a result of this, up to 25% of all
human fetuses are aneuploid, and most of
these result from nondisjunction in oocytes at
meiosis I.
78
• If meiosis goes wrong in male cells, a cell-cycle checkpoint
mechanism is activated, which arrests meiosis and leads to cell
death by apoptosis
• Such checkpoint mechanisms apparently do not operate in female
meiotic cells: if homolog segregation fails to occur normally, the
cells continue through meiosis and produce aneuploid eggs
• The male germ line, on the other hand, is thought to be the main
source of another type of genetic error. Because many more
mitotic cell divisions occur on the way to the production of a
sperm, and each round of DNA replication is liable to error, the
average number of new mutations contributed by fathers is larger
than the number contributed by mothers.

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