Introduction

What do your intestines, the yeast in bread dough,

and a developing frog all have in common? Among
other things, they all have cells that carry out
mitosis, dividing to produce more cells that are
genetically identical to themselves.
Why do these very different organisms and tissues
all need mitosis? Intestinal cells have to be replaced
as they wear out; yeast cells need to reproduce to
keep their population growing; and a tadpole must
make new cells as it grows bigger and more
complex.
What is mitosis?
Mitosis is a type of cell division in which one cell
(the mother) divides to produce two new cells (the
daughters) that are genetically identical to itself. In
the context of the cell cycle, mitosis is the part of the
division process in which the DNA of the cell's
nucleus is split into two equal sets of chromosomes.
The great majority of the cell divisions that happen in
your body involve mitosis. During development and
growth, mitosis populates an organism’s body with
cells, and throughout an organism’s life, it replaces
old, worn-out cells with new ones. For single-celled
eukaryotes like yeast, mitotic divisions are actually a
form of reproduction, adding new individuals to the
population.
In all of these cases, the “goal” of mitosis is to make
sure that each daughter cell gets a perfect, full set of
chromosomes. Cells with too few or too many
chromosomes usually don’t function well: they may
not survive, or they may even cause cancer. So,
when cells undergo mitosis, they don’t just divide
their DNA at random and toss it into piles for the two
daughter cells. Instead, they split up their duplicated
chromosomes in a carefully organized series of
steps.
Phases of mitosis
Mitosis consists of four basic phases: prophase,
metaphase, anaphase, and telophase. Some
textbooks list ve, breaking prophase into an early
phase (called prophase) and a late phase (called
prometaphase). These phases occur in strict
sequential order, and cytokinesis - the process of
dividing the cell contents to make two new cells -
starts in anaphase or telophase.
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 Stages of mitosis: prophase, metaphase, anaphase,
telophase. Cytokinesis typically overlaps with
anaphase and/or telophase.
You can remember the order of the phases with the
famous mnemonic: [Please] Pee on the MAT. But
don’t get too hung up on names – what’s most
important to understand is what’s happening at each
stage, and why it’s important for the division of the
chromosomes.

Late G2 phase. The cell has two centrosomes, each

with two centrioles, and the DNA has been copied.
 At this stage, the DNA is surrounded by an intact
nuclear membrane, and the nucleolus is present in
the nucleus.
Let’s start by looking at a cell right before it begins
mitosis. This cell is in interphase (late G

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phase) and has already copied its DNA, so the
chromosomes in the nucleus each consist of two
connected copies, called sister chromatids. You
can’t see the chromosomes very clearly at this point,
because they are still in their long, stringy,
decondensed form.
This animal cell has also made a copy of its
centrosome, an organelle that will play a key role in
orchestrating mitosis, so there are two centrosomes.
(Plant cells generally don’t have centrosomes with
centrioles, but have a different type of microtubule
organizing center that plays a similar role.)
 Early prophase. The mitotic spindle starts to form,
the chromosomes start to condense, and the
nucleolus disappears.
In early prophase, the cell starts to break down
some structures and build others up, setting the
stage for division of the chromosomes.
• The chromosomes start to condense
(making them easier to pull apart later on).
• The mitotic spindle begins to form. The
spindle is a structure made of microtubules,
strong bers that are part of the cell’s “skeleton.”
Its job is to organize the chromosomes and
move them around during mitosis. The spindle
grows between the centrosomes as they move
apart.
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 • The nucleolus (or nucleoli, plural), a part of
the nucleus where ribosomes are made,
disappears. This is a sign that the nucleus is
getting ready to break down.

Late prophase (prometaphase). The nuclear

envelope breaks down and the chromosomes are
fully condensed.
In late prophase (sometimes also called
prometaphase), the mitotic spindle begins to
capture and organize the chromosomes.
• The chromosomes become even more
condensed, so they are very compact.
• The nuclear envelope breaks down,
releasing the chromosomes.
 • The mitotic spindle grows more, and some
of the microtubules start to “capture”
chromosomes.

Anatomy of the mitotic spindle. Diagram indicating

kinetochore microtubules (bound to kinetochores)
and the aster. The aster is an array of microtubules
that radiates out from the centrosome towards the
cell edge. Diagram also indicates the centromere
region of a chromosome, the narrow "waist" where
the two sister chromatids are most tightly connected,
and the kinetochore, a pad of proteins found at the
centromere.
Microtubules can bind to chromosomes at the
kinetochore, a patch of protein found on the
centromere of each sister chromatid. (Centromeres
are the regions of DNA where the sister chromatids
are most tightly connected.)
Microtubules that bind a chromosome are called
kinetochore microtubules. Microtubules that don’t
bind to kinetochores can grab on to microtubules
from the opposite pole, stabilizing the spindle. More
microtubules extend from each centrosome towards
the edge of the cell, forming a structure called the
aster.

Metaphase. Chromosomes line up at the metaphase

plate, under tension from the mitotic spindle. The
two sister chromatids of each chromosome are
captured by microtubules from opposite spindle
poles.
In metaphase, the spindle has captured all the
chromosomes and lined them up at the middle of the
cell, ready to divide.
• All the chromosomes align at the
metaphase plate (not a physical structure, just
a term for the plane where the chromosomes
line up).
• At this stage, the two kinetochores of each
chromosome should be attached to microtubules
from opposite spindle poles.
Before proceeding to anaphase, the cell will check to
make sure that all the chromosomes are at the
metaphase plate with their kinetochores correctly
attached to microtubules. This is called the spindle
checkpoint and helps ensure that the sister
chromatids will split evenly between the two
daughter cells when they separate in the next step.
If a chromosome is not properly aligned or attached,
the cell will halt division until the problem is xed.
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 Anaphase. The sister chromatids separate from one
another and are pulled towards opposite poles of the
cell. The microtubules that are not attached to
chromosomes push the two poles of the spindle
apart, while the kinetochore microtubules pull the
chromosomes towards the poles.
In anaphase, the sister chromatids separate from
each other and are pulled towards opposite ends of
the cell.
• The protein “glue” that holds the sister
chromatids together is broken down, allowing
them to separate. Each is now its own
chromosome. The chromosomes of each pair
are pulled towards opposite ends of the cell.
 • Microtubules not attached to chromosomes
elongate and push apart, separating the poles
and making the cell longer.
All of these processes are driven by motor
proteins, molecular machines that can “walk” along
microtubule tracks and carry a cargo. In mitosis,
motor proteins carry chromosomes or other
microtubules as they walk.

Telophase. The spindle disappears, a nuclear

membrane re-forms around each set of
chromosomes, and a nucleolus reappears in each
 new nucleus. The chromosomes also start to
decondense.
In telophase, the cell is nearly done dividing, and it
starts to re-establish its normal structures as
cytokinesis (division of the cell contents) takes place.
• The mitotic spindle is broken down into its
building blocks.
• Two new nuclei form, one for each set of
chromosomes. Nuclear membranes and nucleoli
reappear.
• The chromosomes begin to decondense and
return to their “stringy” form.

Cytokinesis in animal and plant cells.

 Cytokinesis in an animal cell: an actin ring around
the middle of the cell pinches inward, creating an
indentation called the cleavage furrow.
Cytokinesis in a plant cell: the cell plate forms down
the middle of the cell, creating a new wall that
partitions it in two.
Cytokinesis, the division of the cytoplasm to form
two new cells, overlaps with the nal stages of
mitosis. It may start in either anaphase or telophase,
depending on the cell, and nishes shortly after
telophase.
In animal cells, cytokinesis is contractile, pinching
the cell in two like a coin purse with a drawstring.
The “drawstring” is a band of laments made of a
protein called actin, and the pinch crease is known
as the cleavage furrow. Plant cells can’t be divided
like this because they have a cell wall and are too
stiff. Instead, a structure called the cell plate forms
down the middle of the cell, splitting it into two
daughter cells separated by a new wall.
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 When division is complete, it produces two daughter
cells. Each daughter cell has a complete set of
chromosomes, identical to that of its sister (and that
of the mother cell). The daughter cells enter the cell
cycle in G1.
When cytokinesis nishes, we end up with two new
cells, each with a complete set of chromosomes
identical to those of the mother cell. The daughter
cells can now begin their own cellular “lives,” and –
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depending on what they decide to be when they
grow up – may undergo mitosis themselves,
repeating the cycle.