Important Terms

Cell cycle - A repeated process in which cells copy their contents and then
divide into two. Occurs in cells as a means of reproduction.

Diploid number - The total number of chromosomes present in a cell.

Gametes - Cells with the haploid number of chromosomes. Gametes are

created out of germ cells. The sperm and eggs involved in sexual
reproduction are gametes.

Germ cell - Cells that lead to the production of gametes. Produced by

meiosis.

Haploid number - The number of unique chromosomes or homologous

pairs in a cell. Half the diploid number.

Homologous pair - Refers to two similar chromosomes in a diploid cell.

One chromosome is derived from the father gamete cell and the other from
the mother gamete.

Meiosis - A type of cellular reproduction that results in the formation of

four haploid cells from one diploid cell. Contains two cellular divisions that
follow only one round of DNA replication. The type of reproduction that
produces germ cells.

Mitosis - A type of cellular reproduction that results in the formation of two

identical diploid cells from one diploid cell. Contains one cellular division
following a round of DNA replication. The type of reproduction that produces
somatic cells.

Sister chromatid - Refers to the copy of a chromosomes that results from

DNA replication and is still closely linked to its original.

Somatic cell - Any plant or animal cell that is not a germ cell. The class of
cell formed during mitosis.
Components of the Cell Cycle

The Cell Cycle

The two main events of cellular reproduction are the copying of cellular
components and the cleavage of the cell. These two events, copying and
cleaving, represent the two larger phases of the cell cycle, interphase and
Mitosis. Mitosis is the part of the cell cycle when the cell prepares for and
completes cell division. During interphase, appropriate cellular components
are copied. Interphase is also a time of checkpoints to make sure that the
cell is ready to proceed into mitosis. Both of these two phases have further
sub-divisions. Since the cell cycle is a "cycle" it has no distinct beginning or
ending. Cells are continually entering and exiting the various phases of the
cycle.

Figure %: Interphase and Mitosis

Interphase

We will begin our discussion of the events that take place during interphase
with those that occur immediately after a cell has successfully divided during
mitosis. This phase is called G1.

G1 phase

G1 is an intermediate phase occupying the time between the end of cell

division in mitosis and the beginning of DNA replication during S phase.
During this time, the cell grows in preparation for DNA replication, and
certain intracellular components, such as the centrosomes undergo
replication. Before a cell begins DNA replication, it must ensure that it is
biologically ready to take on such a process. G1 is the phase when this
cellular monitoring takes place.

During G1, the cell reviews the cellular environment and the cell size to
ensure that the conditions are appropriate to support DNA replication. Not
until the cell is ready does it leave G1. If all is not ready to undergo DNA
replication, cells can pause during G1 and enter a phase called G0.
Depending on a cell's preparedness to continue in the cell cycle, G0 can last
days, weeks, or even years. When the cell has reached an appropriate size
and is in a supportive environment for DNA replication, it will exit either G1
or G0 and enter the next phase of interphase called S phase.

S phase

S phase, or synthesis, is the phase of the cell cycle when DNA packaged into
chromosomes is replicated. This event is an essential aspect of the cell cycle
because replication allows for each cell created by cell division to have the
same genetic make-up. During S phase a number of events additional to
chromosome replication take place. Cell growth continues through S phase,
as does the rate of synthesis of a number of proteins and enzymes that are
involved in DNA synthesis. Once DNA replication is complete the cell contains
twice its normal number of chromosomes and becomes ready to enter the
phase called G2.

G2 phase

Similar to G1, G2 is an intermediate phase, a time for the cell to ensure that
it is ready to proceed in the cell cycle. Occurring between the end of DNA
replication in S phase and the beginning of cell division in mitosis, G2 can be
thought of as a safety gap during which a cell can check to make sure that
the entirety of its DNA and other intracellular components have been
properly duplicated. In addition to acting as a checkpoint along the cell cycle,
G2 also represents the cell's final chance to grow before it is split into two
independent cells during mitosis.

Cell Cycle Summary

Interphase is made up of three distinct phases: G1, S phase, and G2. The G1
and G2 phases serve as checkpoints for the cell to make sure that it is ready
to proceed in the cell cycle. If it is not, the cell will use this time to make
proper adjustments that can include cell growth, correction or completion of
DNA synthesis, and duplication of intracellular components. S phase involves
the replication of chromosomes. All three stages of interphase involve
continued cell growth and an increase in the concentration of proteins found
in the cell.

Prophase

As we discussed in cell cycle, before cells are allowed to enter M phase they
must meet certain cellular requirements. Among these requirements are
appropriate cell size and cellular environment. Following DNA replication in S
phase, cells contain twice their normal number of chromosomes. Because
cells that undergo mitosis are diploid, their number of chromosomes can be
represented as 2N, where N equals the number of distinct chromosomes in
the cell. Cells about to enter M phase, which have passed through S phase
and replicated their DNA, have 4N chromosomes.
The first phase of mitosis within M phase is called prophase. It follows G2,
the final phase of interphase. A cell entering M phase manifests a number of
physicsl signs. Among these are condensation, or thickening, of
chromosomes. Chromosome condensation is visible through a microscope
and is required for subsequent chromosome separation during later stages of
mitosis. Another physical characteristic of cells beginning mitosis is the
sprouting of microtubules from replicated centrosomes. Microtubles are
protein filaments on which chromosomes migrate during mitosis.

Prophase

Figure 1: Prophase

As we discussed, prophase is marked by very thick and dense chromosomes.

At this phase, the chromosomes are still enclosed in the cell nucleus within
the nuclear envelope. The chromosomes also contain a centromere, which is
necessary in later phases for attachment to microtubules for migration. Late
in prophase, kinetochores assemble on the centromeres. Specialized
microtubules, called kinetochore microtubules later attach to these sites.
Duplicated centrosomes, which are the organizing centers of microtubules,
begin to separate towards opposite poles of the cell. The network of
cytoskeletal components begins to break down and the mitotic spindle forms.
The mitotic spindle is an arrangement of microtubules that is responsible for
aligning duplicated chromosomes in later phases.

Metaphase and Anaphase

The next two major events that take place in mitosis are the alignment of
chromosomes at the center of the cell and the subsequent separation of
sister chromatids to opposite mitotic spindle poles. These two events occur
in metaphase and anaphase, respectively. In this section we will review the
events of both of these phases.

Metaphase
Figure 2: Metaphase
The centrosomes have aligned at opposite ends, or poles of the cell and
chromosomes are being moved toward the center of the cell. Metaphase is
marked by the alignment of chromosomes at the center of the cell, half way
between each of the mitoic spindle poles. Movement is mediated by the
kinetochore microtubles, which push and pull on the chromosomes to align
them into what is called the metaphase plate. Chromosomes on the
metaphase plate are held there tightly by pushing and pulling forces from
the microtubules.
Microtubule structure allows them to be dynamic molecules. The subunit of
microtubules is called tubulin and it is constantly added and removed from
the ends of microtubules leading to a state of treadmilling. The
chromosomes are held tightly by these forces constantly pushing and pulling
on them.
Metaphase can occupy a large portion of the total time of mitosis because
chromosome alignment at the center of the cell on the metaphase plate acts
as a checkpoint for progression into the next phase, anaphase. Cells can
arrest in metaphase for days until the chromosomes are properly aligned and
the cell enters anaphase.
Anaphase
Figure 3: Anaphase
Entrance into anaphase is triggered by the inactivation of M phase-promoting
factor that follows mitotic cyclin degradation. During anaphase, the
kinetochore microtubules retract, increasing the seperation of the sister
chromatids as they are moved further toward the opposite spindle poles.
Anaphase can be broken into two distinct phases. In the first phase,
called anaphase A, chromosomes move poleward, away from the metaphase
plate with the retraction of the microtubules. This movement occurs at
approximately 2 micrometers per minute (the entire length of a cell is
between 10 and 30 micrometers). In the second phase, anaphase B, the
mitotic poles marked by the centrosomes themselves separate by the
elongation of a specific type of non-kinetochore microtubule, called a polar
microtubule. The extent of the separation of the poles varies from species to
species. The entire duration of anaphase is relatively short, usually only
lasting a few minutes.

Telophase and Cytokinesis

The final two events of M phase are the re-forming of the nuclear envelope
around the separated sister chromatids and the cleavage of the cell. These
events occur in telophase and cytokinesis, respectively. In this section we
will review the events that comprise these final phases of M phase.
Telophase

Figure 4: Telophase

Telophase is technically the final stage of mitosis. Its name derives from the
latin word telos which means end. During this phase, the sister chromatids
reach opposite poles. The small nuclear vesicles in the cell begin to re-form
around the group of chromosomes at each end. As the nuclear envelope re-
forms by associating with the chromosomes, two nuclei are created in the
one cell. Telophase is also marked by the dissolution of the kinetochore
microtubules and the continued elongation of the polar microtubules. As the
nuclear envelopes re-form, the chromosomes begin to decondense and
become more diffuse.

Cytokinesis
Figure 5 : Cytokinesis

Cytokinesis is the process in which the cell actually divides into two. With the
two nuclei already at opposite poles of the cell, the cell cytoplasm separates,
and the cell pinches in the middle, ultimately leading to cleavage. In most
cells, the mitotic spindle determines the site where the cell will begin to
invaginate and split. The first signs of this puckering are usually visible
sometime during anaphase.
Earlier we mentioned that in prophase, the cell's cytoskeleton becomes
disassembled. The disassembled cytoskeletal filaments are used in a
different way during cytokinesis. Cleavage occurs by the contraction of a thin
ring of actin filaments that form the contractile ring. The contractile ring
defines the cleavage line for the cell. If the ring is not positioned at the
center of the cell, an asymmetrical division takes place. The ring contracts
and eventually pinches the cell until it separates into two independent
daughter cells. In higher order plants, the cytokinesis process is slightly
different because the cytoplasm splits with the formation of the cell wall.