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Meiosis

How meiosis reduces chromosome number by half: crossing over, meiosis I, meiosis II, and genetic
variation.

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Introduction

Mitosis is used for almost all of your body’s cell division needs. It adds new cells during development
and replaces old and worn-out cells throughout your life. The goal of mitosis is to produce daughter cells
that are genetically identical to their mothers, with not a single chromosome more or less.

Meiosis, on the other hand, is used for just one purpose in the human body: the production of
gametes—sex cells, or sperm and eggs. Its goal is to make daughter cells with exactly half as many
chromosomes as the starting cell.

To put that another way, meiosis in humans is a division process that takes us from a diploid cell—one
with two sets of chromosomes—to haploid cells—ones with a single set of chromosomes. In humans,
the haploid cells made in meiosis are sperm and eggs. When a sperm and an egg join in fertilization, the
two haploid sets of chromosomes form a complete diploid set: a new genome.

Phases of meiosis

In many ways, meiosis is a lot like mitosis. The cell goes through similar stages and uses similar strategies
to organize and separate chromosomes. In meiosis, however, the cell has a more complex task. It still
needs to separate sister chromatids (the two halves of a duplicated chromosome), as in mitosis. But it
must also separate homologous chromosomes, the similar but nonidentical chromosome pairs an
organism receives from its two parents.

These goals are accomplished in meiosis using a two-step division process. Homologue pairs separate
during a first round of cell division, called meiosis I. Sister chromatids separate during a second round,
called meiosis II.

Since cell division occurs twice during meiosis, one starting cell can produce four gametes (eggs or
sperm). In each round of division, cells go through four stages: prophase, metaphase, anaphase, and
telophase.

Meiosis I

Before entering meiosis I, a cell must first go through interphase. As in mitosis, the cell grows during G_1

phase, copies all of its chromosomes during S phase, and prepares for division during G_2

phase.
During prophase I, differences from mitosis begin to appear. As in mitosis, the chromosomes begin to
condense, but in meiosis I, they also pair up. Each chromosome carefully aligns with its homologue
partner so that the two match up at corresponding positions along their full length.

For instance, in the image below, the letters A, B, and C represent genes found at particular spots on the
chromosome, with capital and lowercase letters for different forms, or alleles, of each gene. The DNA is
broken at the same spot on each homologue—here, between genes B and C—and reconnected in a
criss-cross pattern so that the homologues exchange part of their DNA.

Image of crossing over. Two homologous chromosomes carry different versions of three genes. One has
the A, B, and C versions, while the other has the a, b, and c versions. A crossover event in which two
chromatids—one from each homologue—exchange fragments swaps the C and c genes. Now, each
homologue has two dissimilar chromatids.

One has A, B, C on one chromatid and A, B, c on the other chromatid.

The other homologue has a, b, c on one chromatid and a, b, C on the other chromatid.

Image of crossing over. Two homologous chromosomes carry different versions of three genes. One has
the A, B, and C versions, while the other has the a, b, and c versions. A crossover event in which two
chromatids—one from each homologue—exchange fragments swaps the C and c genes. Now, each
homologue has two dissimilar chromatids.

One has A, B, C on one chromatid and A, B, c on the other chromatid.

The other homologue has a, b, c on one chromatid and a, b, C on the other chromatid.

Image credit: based on "The process of meiosis: Figure 2" by OpenStax College, Biology, CC BY 3.0

This process, in which homologous chromosomes trade parts, is called crossing over. It's helped along by
a protein structure called the synaptonemal complex that holds the homologues together. The
chromosomes would actually be positioned one on top of the other—as in the image below—
throughout crossing over; they're only shown side-by-side in the image above so that it's easier to see
the exchange of genetic material.

Image of two homologous chromosomes, positioned one on top of the other and held together by the
synaptonemal complex.

Image of two homologous chromosomes, positioned one on top of the other and held together by the
synaptonemal complex.

Image credit: based on "The process of meiosis: Figure 1" by OpenStax College, Biology, CC BY 3.0

You can see crossovers under a microscope as chiasmata, cross-shaped structures where homologues
are linked together. Chiasmata keep the homologues connected to each other after the synaptonemal
complex breaks down, so each homologous pair needs at least one. It's common for multiple crossovers
(up to 2525!) to take place for each homologue pair ^1
1

The spots where crossovers happen are more or less random, leading to the formation of new,
"remixed" chromosomes with unique combinations of alleles.

After crossing over, the spindle begins to capture chromosomes and move them towards the center of
the cell (metaphase plate). This may seem familiar from mitosis, but there is a twist. Each chromosome
attaches to microtubules from just one pole of the spindle, and the two homologues of a pair bind to
microtubules from opposite poles. So, during metaphase I, homologue pairs—not individual
chromosomes—line up at the metaphase plate for separation.

The phases of meiosis I.

Prophase I: The starting cell is diploid, 2n = 4. Homologous chromosomes pair up and exchange
fragments in the process of crossing over.

Metaphase I: Homologue pairs line up at the metaphase plate.

Anaphase I: Homologues separate to opposite ends of the cell. Sister chromatids stay together.

Telophase I: Newly forming cells are haploid, n = 2. Each chromosome still has two sister chromatids, but
the chromatids of each chromosome are no longer identical to each other.

The phases of meiosis I.

Prophase I: The starting cell is diploid, 2n = 4. Homologous chromosomes pair up and exchange
fragments in the process of crossing over.

Metaphase I: Homologue pairs line up at the metaphase plate.

Anaphase I: Homologues separate to opposite ends of the cell. Sister chromatids stay together.

Telophase I: Newly forming cells are haploid, n = 2. Each chromosome still has two sister chromatids, but
the chromatids of each chromosome are no longer identical to each other.

When the homologous pairs line up at the metaphase plate, the orientation of each pair is random. For
instance, in the diagram above, the pink version of the big chromosome and the purple version of the
little chromosome happen to be positioned towards the same pole and go into the same cell. But the
orientation could have equally well been flipped, so that both purple chromosomes went into the cell
together. This allows for the formation of gametes with different sets of homologues. [Can you show me
what you mean?]
Diagram showing the relationship between chromosome configuration at meiosis I and homologue
segregation to gametes. The diagram depicts a simplified case in which an organism only has 2n = 4
chromosomes. In this case, four different types of gametes may be produced, depending on whether
the maternal homologues are positioned on the same side or on opposite sides of the metaphase plate.

Diagram showing the relationship between chromosome configuration at meiosis I and homologue
segregation to gametes. The diagram depicts a simplified case in which an organism only has 2n = 4
chromosomes. In this case, four different types of gametes may be produced, depending on whether
the maternal homologues are positioned on the same side or on opposite sides of the metaphase plate.

In anaphase I, the homologues are pulled apart and move apart to opposite ends of the cell. The sister
chromatids of each chromosome, however, remain attached to one another and don't come apart.

Finally, in telophase I, the chromosomes arrive at opposite poles of the cell. In some organisms, the
nuclear membrane re-forms and the chromosomes decondense, although in others, this step is
skipped—since cells will soon go through another round of division, meiosis II^{2,3}

2,3

. Cytokinesis usually occurs at the same time as telophase I, forming two haploid daughter cells.

Meiosis II

Cells move from meiosis I to meiosis II without copying their DNA. Meiosis II is a shorter and simpler
process than meiosis I, and you may find it helpful to think of meiosis II as “mitosis for haploid cells."

The cells that enter meiosis II are the ones made in meiosis I. These cells are haploid—have just one
chromosome from each homologue pair—but their chromosomes still consist of two sister chromatids.
In meiosis II, the sister chromatids separate, making haploid cells with non-duplicated chromosomes.

Phases of meiosis II

Prophase II: Starting cells are the haploid cells made in meiosis I. Chromosomes condense.

Metaphase II: Chromosomes line up at the metaphase plate.

Anaphase II: Sister chromatids separate to opposite ends of the cell.

Telophase II: Newly forming gametes are haploid, and each chromosome now has just one chromatid.

Phases of meiosis II

Prophase II: Starting cells are the haploid cells made in meiosis I. Chromosomes condense.

Metaphase II: Chromosomes line up at the metaphase plate.


Anaphase II: Sister chromatids separate to opposite ends of the cell.

Telophase II: Newly forming gametes are haploid, and each chromosome now has just one chromatid.

During prophase II, chromosomes condense and the nuclear envelope breaks down, if needed. The
centrosomes move apart, the spindle forms between them, and the spindle microtubules begin to
capture chromosomes. [When did the centrosomes duplicate?]

^3

^5

The two sister chromatids of each chromosome are captured by microtubules from opposite spindle
poles. In metaphase II, the chromosomes line up individually along the metaphase plate. In anaphase II,
the sister chromatids separate and are pulled towards opposite poles of the cell.

In telophase II, nuclear membranes form around each set of chromosomes, and the chromosomes
decondense. Cytokinesis splits the chromosome sets into new cells, forming the final products of
meiosis: four haploid cells in which each chromosome has just one chromatid. In humans, the products
of meiosis are sperm or egg cells. [Does meiosis always produce four gametes?]

^2

How meiosis "mixes and matches" genes

The gametes produced in meiosis are all haploid, but they're not genetically identical. For example, take
a look the meiosis II diagram above, which shows the products of meiosis for a cell with 2n = 42n=4
chromosomes. Each gamete has a unique "sample" of the genetic material present in the starting cell.

As it turns out, there are many more potential gamete types than just the four shown in the diagram,
even for a cell with only four chromosomes. The two main reasons we can get many genetically different
gametes are:

Crossing over. The points where homologues cross over and exchange genetic material are chosen more
or less at random, and they will be different in each cell that goes through meiosis. If meiosis happens
many times, as in humans, crossovers will happen at many different points.

Random orientation of homologue pairs. The random orientation of homologue pairs in metaphase I
allows for the production of gametes with many different assortments of homologous chromosomes.

In a human cell, the random orientation of homologue pairs alone allows for over 88
\text{million}million different types of possible gametes^7

7
. When we layer crossing over on top of this, the number of genetically different gametes that you—or
any other person—can make is effectively infinite. [How do you get that number?]

23

2^2 = 4

232^{23}=8,

388,608

Diagram showing the relationship between chromosome configuration at meiosis I and homologue
segregation to gametes. The diagram depicts a simplified case in which an organism only has 2n = 4
chromosomes. In this case, four different types of gametes may be produced, depending on whether
the maternal homologues are positioned on the same side or on opposite sides of the metaphase plate.

Diagram showing the relationship between chromosome configuration at meiosis I and homologue
segregation to gametes. The diagram depicts a simplified case in which an organism only has 2n = 4
chromosomes. In this case, four different types of gametes may be produced, depending on whether
the maternal homologues are positioned on the same side or on opposite sides of the metaphase plate.

Check out the video on variation in a species to learn how genetic diversity generated by meiosis (and
fertilization) is important in evolution and helps populations survive.
The Cell Cycle, Mitosis and Meiosis

Topic related resources

The cell cycle

Actively dividing eukaryote cells pass through a series of stages known collectively as the cell cycle: two
gap phases (G1 and G2); an S (for synthesis) phase, in which the genetic material is duplicated; and an M
phase, in which mitosis partitions the genetic material and the cell divides.

22-Cell-cycle.gif

G1 phase. Metabolic changes prepare the cell for division. At a certain point - the restriction point - the
cell is committed to division and moves into the S phase.

S phase. DNA synthesis replicates the genetic material. Each chromosome now consists of two sister
chromatids.

G2 phase. Metabolic changes assemble the cytoplasmic materials necessary for mitosis and cytokinesis.

M phase. A nuclear division (mitosis) followed by a cell division (cytokinesis).

The period between mitotic divisions - that is, G1, S and G2 - is known as interphase.

Mitosis

Mitosis is a form of eukaryotic cell division that produces two daughter cells with the same genetic
component as the parent cell. Chromosomes replicated during the S phase are divided in such a way as
to ensure that each daughter cell receives a copy of every chromosome. In actively dividing animal cells,
the whole process takes about one hour.

The replicated chromosomes are attached to a 'mitotic apparatus' that aligns them and then separates
the sister chromatids to produce an even partitioning of the genetic material. This separation of the
genetic material in a mitotic nuclear division (or karyokinesis) is followed by a separation of the cell
cytoplasm in a cellular division (or cytokinesis) to produce two daughter cells.

In some single-celled organisms mitosis forms the basis of asexual reproduction. In diploid multicellular
organisms sexual reproduction involves the fusion of two haploid gametes to produce a diploid zygote.
Mitotic divisions of the zygote and daughter cells are then responsible for the subsequent growth and
development of the organism. In the adult organism, mitosis plays a role in cell replacement, wound
healing and tumour formation.
Mitosis, although a continuous process, is conventionally divided into five stages: prophase,
prometaphase, metaphase, anaphase and telophase.

38 mitotis phases.jpg

The phases of mitosis

Prophase

Prophase occupies over half of mitosis. The nuclear membrane breaks down to form a number of small
vesicles and the nucleolus disintegrates. A structure known as the centrosome duplicates itself to form
two daughter centrosomes that migrate to opposite ends of the cell. The centrosomes organise the
production of microtubules that form the spindle fibres that constitute the mitotic spindle. The
chromosomes condense into compact structures. Each replicated chromosome can now be seen to
consist of two identical chromatids (or sister chromatids) held together by a structure known as the
centromere.

Prometaphase

The chromosomes, led by their centromeres, migrate to the equatorial plane in the mid-line of the cell -
at right-angles to the axis formed by the centrosomes. This region of the mitotic spindle is known as the
metaphase plate. The spindle fibres bind to a structure associated with the centromere of each
chromosome called a kinetochore. Individual spindle fibres bind to a kinetochore structure on each side
of the centromere. The chromosomes continue to condense.

Metaphase

The chromosomes align themselves along the metaphase plate of the spindle apparatus.

Anaphase

The shortest stage of mitosis. The centromeres divide, and the sister chromatids of each chromosome
are pulled apart - or 'disjoin' - and move to the opposite ends of the cell, pulled by spindle fibres
attached to the kinetochore regions. The separated sister chromatids are now referred to as daughter
chromosomes. (It is the alignment and separation in metaphase and anaphase that is important in
ensuring that each daughter cell receives a copy of every chromosome.)

Telophase

The final stage of mitosis, and a reversal of many of the processes observed during prophase. The
nuclear membrane reforms around the chromosomes grouped at either pole of the cell, the
chromosomes uncoil and become diffuse, and the spindle fibres disappear.

Cytokinesis

The final cellular division to form two new cells. In plants a cell plate forms along the line of the
metaphase plate; in animals there is a constriction of the cytoplasm. The cell then enters interphase -
the interval between mitotic divisions.

Meiosis

Meiosis is the form of eukaryotic cell division that produces haploid sex cells or gametes (which contain
a single copy of each chromosome) from diploid cells (which contain two copies of each chromosome).
The process takes the form of one DNA replication followed by two successive nuclear and cellular
divisions (Meiosis I and Meiosis II). As in mitosis, meiosis is preceded by a process of DNA replication
that converts each chromosome into two sister chromatids.

Meiosis I

Meiosis I separates the pairs of homologous chromosomes.

39-meiosis_I_males.gif

In Meiosis I a special cell division reduces the cell from diploid to haploid.

Prophase I

The homologous chromosomes pair and exchange DNA to form recombinant chromosomes. Prophase I
is divided into five phases:
Leptotene: chromosomes start to condense.

Zygotene: homologous chromosomes become closely associated (synapsis) to form pairs of


chromosomes (bivalents) consisting of four chromatids (tetrads).

Pachytene: crossing over between pairs of homologous chromosomes to form chiasmata (sing.
chiasma).

Diplotene: homologous chromosomes start to separate but remain attached by chiasmata.

Diakinesis: homologous chromosomes continue to separate, and chiasmata move to the ends of the
chromosomes.

Prometaphase I

Spindle apparatus formed, and chromosomes attached to spindle fibres by kinetochores.

Metaphase I

Homologous pairs of chromosomes (bivalents) arranged as a double row along the metaphase plate. The
arrangement of the paired chromosomes with respect to the poles of the spindle apparatus is random
along the metaphase plate. (This is a source of genetic variation through random assortment, as the
paternal and maternal chromosomes in a homologous pair are similar but not identical. The number of
possible arrangements is 2n, where n is the number of chromosomes in a haploid set. Human beings
have 23 different chromosomes, so the number of possible combinations is 223, which is over 8 million.)

Anaphase I

The homologous chromosomes in each bivalent are separated and move to the opposite poles of the
cell

Telophase I

The chromosomes become diffuse and the nuclear membrane reforms.

Cytokinesis

The final cellular division to form two new cells, followed by Meiosis II. Meiosis I is a reduction division:
the original diploid cell had two copies of each chromosome; the newly formed haploid cells have one
copy of each chromosome.

Meiosis II
Meiosis II separates each chromosome into two chromatids.

40-meiosis_II_males.gif

The events of Meiosis II are analogous to those of a mitotic division, although the number of
chromosomes involved has been halved.

Meiosis generates genetic diversity through:

the exchange of genetic material between homologous chromosomes during Meiosis I

the random alignment of maternal and paternal chromosomes in Meiosis I

the random alignment of the sister chromatids at Meiosis II

Meiosis in females

41-meiosis_I_females.gif

42-meiosis_II_females.gif

Topic related resources