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.
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 G1 phase, copies all of its chromosomes during S phase, and prepares
for division during G2 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.

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.
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 25!) to take place for
each homologue pair.
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.
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.
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. 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.
 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.
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.
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=42, n, equals, 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 8
million different types of possible gametes. 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.
GUIDE QUESTIONS
1. What is meiosis?
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2. What is the difference between a haploid cell and a diploid cell?

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3. Describe the steps of Meiosis I and Meiosis II.

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4. What is the result of crossing-over?

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5. State the phase where each of the following processes occurs: (a) sister chromatids separate,
(b) homologous chromosomes form pairs, (c) two haploid cells form.
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6. At the end of meiosis II, each of the haploid sex cells has only half the number of
chromosomes as the original diploid cell. Why is this important?
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