Reading 4.

2 Cell cycle

Table of Contents – ctrl+click to jump to a topic

Introduction.....................................................................................................2
Learning objectives.......................................................................................2
Videos to Watch............................................................................................3
Text to Read....................................................................................................4
DNA organization in cells..............................................................................4
Cell cycle.......................................................................................................7
Interphase..................................................................................................8
Mitotic phase..............................................................................................9
Prokaryotic cell division..............................................................................13

Immunofluorescent light micrograph of a cell (center) during the metaphase stage of mitosis
(cell division). It is surrounded by interphase (resting) cells. During metaphase, the DNA-
containing chromosomes (yellow/orange) attach to the microtubules (green) and form a line
(the metaphase plate) on the microtubule spindle. The microtubules pull the chromosomes
apart towards either end of the cell. The cell will eventually split in two, forming genetically-
identical daughter cells. Immunofluorescence uses antibodies to attach fluorescent dyes to
specific cell tissues. Credit: Dr. Paul Andrews, University of Dundee / Science Source
Reading 4.2 Cell cycle

Introduction
Cell division, a fundamental process in biology, has its roots in the earliest
forms of life. It ensures the continuity of life by allowing organisms to grow,
repair tissues, and reproduce. While the basic principles of cell division are
conserved across different organisms, the specific mechanisms and
outcomes vary significantly. Prokaryotes, the simplest form of life, divide
through a relatively simple process called binary fission. In contrast,
eukaryotic cells, which possess a nucleus and other membrane-bound
organelles, undergo a more complex process known as mitosis. This process
involves precise duplication of genetic material and the division of the cell
into two identical daughter cells.
Cell division is essential for a wide range of biological processes. For
example, in bacteria, it allows for rapid population growth, while in
multicellular organisms, it is crucial for growth, development, and tissue
repair. In plants, cell division is involved in the formation of new organs and
the generation of gametes for sexual reproduction. In animals, it plays a vital
role in embryonic development, tissue regeneration, and the maintenance of
homeostasis.

Learning objectives
After reading completing this reading, you will be able to:
● LO 4.2.1 - Describe the events that occur in the cell cycle
● LO 4.2.2 - Explain how mitosis results in the transmission of chromosomes from
one generation to the next
Reading 4.2 Cell cycle

Videos to Watch

How DNA is packaged

(DNA Learning Center)

Binary fission in bacteria

(Microbial Zoo)

The cell cycle

(Nucleus Biology)
Reading 4.2 Cell cycle

Text to Read
Adapted from OpenStax Biology for AP Courses. Excerpts from chapter 10. Access
for free at openstax.org

DNA organization in cells

Before discussing the steps a cell must undertake to replicate, a deeper
understanding of the structure and function of a cell’s genetic information is
necessary. A cell’s DNA, packaged as a double-stranded DNA molecule, is
called its genome. Recall that in prokaryotes, the genome is composed of a
single, double-stranded DNA molecule in the form of a loop or circle (Figure
1). The region in the cell containing this genetic material is called a nucleoid.
Some prokaryotes also have smaller loops of DNA called plasmids that are
not essential for normal growth. These will be discussed in later readings.

Figure 1 Bacteria have a single circular chromosome that is located in the cytoplasm in a
structure called the nucleoid. Bacteria also contain smaller circular DNA molecules called
plasmids. Credit: The University of Waikato Te Whare Wānanga o Waikato

In eukaryotes, the genome consists of several double-stranded linear DNA

molecules (Figure 2). Each species of eukaryotes has a characteristic number
of chromosomes in the nuclei of its cells. Human body cells have 46
chromosomes, while human gametes (sperm or eggs) have 23
chromosomes each. A typical body cell, or somatic cell, contains two
matched sets of chromosomes, a configuration known as diploid. One set of
each chromosome comes from each parent. The letter n is used to represent
a single set of chromosomes; therefore, a diploid organism is designated 2n.
Human cells that contain one set of chromosomes are called gametes, or sex
cells; these are eggs and sperm, and are designated 1n, or haploid.
Reading 4.2 Cell cycle

Figure 2 There are 23 pairs of homologous chromosomes in a female human somatic cell.
The condensed chromosomes are viewed within the nucleus (top), removed from a cell in
mitosis and spread out on a slide (right), and artificially arranged according to length (left);
an arrangement like this is called a karyotype. In this image, the chromosomes were
exposed to fluorescent stains for differentiation of the different chromosomes. A method of
staining called “chromosome painting” employs fluorescent dyes that highlight
chromosomes in different colors. (credit: National Human Genome Project/NIH)

Matched pairs of chromosomes in a diploid organism are called homologous

(“same knowledge”) chromosomes. Homologous chromosomes are the same
length and have specific nucleotide segments called genes in exactly the
same location, or locus. Genes, the functional units of chromosomes,
determine specific characteristics by coding for specific proteins. Traits are
the variations of those characteristics. For example, hair color is a
characteristic with traits that are blonde, brown, or black.

Each copy of a homologous pair of chromosomes originates from a different

parent; therefore, the genes themselves are not identical. The variation of
individuals within a species is due to the specific combination of the genes
inherited from both parents. Even a slightly altered sequence of nucleotides
within a gene can result in an alternative trait. Minor variations of traits, such
as blood type, eye color, and handedness, contribute to the natural variation
found within a species. However, if the entire DNA sequence from any pair of
human homologous chromosomes is compared, the difference is less than
one percent.
Reading 4.2 Cell cycle

The sex chromosomes, X and Y, are the single exception to the rule of
homologous chromosome uniformity: Other than a small amount of
homology that is necessary to accurately produce gametes, the genes found
on the X and Y chromosomes are different, and, unlike autosomes (non-sex
chromosomes) the two sex chromosomes are different in shape and size, as
well (Figure 3). In humans, a female will inherit one X chromosome from
each parent, and thus carry two X chromosomes in each of her cells; a male
will inherit one X chromosome from his biological mother and one Y
chromosome from his biological father.

Figure 3 A colorized scanning electron micrograph of chromosomes X (left) and Y (right).

Note that unlike homologous autosomal chromosomes, the sex chromosomes are different
sizes.

If the DNA from all 46 chromosomes in a human cell nucleus was laid out end
to end, it would measure approximately two meters; however, its diameter
would be only 2 nm. Considering that the size of a typical human cell is
about 10 µm (100,000 cells lined up to equal one meter), DNA must be
tightly packaged to fit in the cell’s nucleus. At the same time, it must also be
readily accessible for the genes to be expressed. During some stages of the
cell cycle, the long strands of DNA are condensed into compact
chromosomes. There are a number of ways that chromosomes are
compacted.

To achieve its compact state, DNA wraps around histone proteins, forming
structures known as nucleosomes. A nucleosome consists of a DNA
segment wound around a core of eight histone proteins. These nucleosomes
are further compacted into chromatin, a complex of DNA and proteins. This
intricate packaging of DNA ensures efficient storage and protection of
genetic information within the cell nucleus.
Reading 4.2 Cell cycle

Figure 4 Eukaryotic DNA is tightly packaged into chromosomes.

Before a cell divides, each chromosome is duplicated, resulting in two

identical copies called sister chromatids. These sister chromatids are held
together at a specific region called the centromere (Figure 5). Once the cell
divides, each sister chromatid becomes an independent chromosome,
ensuring that each daughter cell receives a complete set of genetic
information.

Figure 5 DNA replication creates identical sister chromatids, joined at a centromere. Credit:
BioNinja

Cell cycle
The cell cycle is an ordered series of events involving cell growth and cell
division that produces two new daughter cells. Cells on the path to cell
division proceed through a series of precisely timed and carefully regulated
stages of growth, DNA replication, and division that produces two identical
(clone) cells. The cell cycle has two major phases: interphase and the mitotic
Reading 4.2 Cell cycle

phase (Figure 6). During interphase, the cell grows and DNA is replicated.
During the mitotic phase, the replicated DNA and cytoplasmic contents are
separated, and the cell divides.

Figure 6 The cell cycle consists of interphase and the mitotic phase. During interphase, the
cell grows and the nuclear DNA is duplicated. Interphase is followed by the mitotic phase.
During the mitotic phase, the duplicated chromosomes are segregated and distributed into
daughter nuclei. The cytoplasm is usually divided as well, resulting in two daughter cells.

Interphase
During interphase, the cell undergoes normal growth processes while also
preparing for cell division. Chromatin is in its least condensed state and
appears loosely distributed throughout the nucleus. The three stages of
interphase are called G1, S, and G2.

The first stage of interphase is called the G1 phase (first gap) because, from
a microscopic aspect, little change is visible. However, during the G1 stage,
the cell is quite active at the biochemical level. The cell is accumulating the
building blocks of chromosomal DNA and the associated proteins as well as
accumulating sufficient energy reserves to complete the task of replicating
each chromosome in the nucleus.

The S phase, or synthesis phase, is a critical stage in the cell cycle where
DNA replication occurs. During this phase, each chromosome is duplicated,
resulting in two identical sister chromatids. This ensures that each daughter
cell receives a complete set of genetic information. In animal cells, the
centrosome, which contains a pair of centrioles, duplicates during the S
phase. These centrosomes will later play a crucial role in organizing the
Reading 4.2 Cell cycle

mitotic spindle, a structure responsible for separating the sister chromatids

during cell division. While the S phase is essential for cell division in both
animal and plant cells, plant cells typically lack centrioles. Instead, they rely
on other mechanisms to organize the mitotic spindle.

In the G2 phase, the cell replenishes its energy stores and synthesizes
proteins necessary for chromosome manipulation. Some cell organelles are
duplicated, and the cytoskeleton is dismantled to provide resources for the
mitotic phase. There may be additional cell growth during G 2. The final
preparations for the mitotic phase must be completed before the cell is able
to enter the first stage of mitosis.

Mitotic phase
The mitotic (M) phase is a multistep process during which the duplicated
chromosomes are aligned, separated, and moved into two new, identical
daughter cells. The first portion of the mitotic phase is called karyokinesis,
or nuclear division. The second portion of the mitotic phase, called
cytokinesis, is the physical separation of the cytoplasmic components into
the two daughter cells.

Karyokinesis, also known as mitosis, is divided into a series of phases—

prophase, prometaphase, metaphase, anaphase, and telophase—that result
in the division of the cell nucleus.
Reading 4.2 Cell cycle

Figure 7 Karyokinesis (or mitosis) is divided into five stages—prophase, prometaphase,

metaphase, anaphase, and telophase. The pictures at the bottom were taken by
fluorescence microscopy (hence, the black background) of cells artificially stained by
fluorescent dyes: blue fluorescence indicates DNA (chromosomes) and green fluorescence
indicates microtubules (spindle apparatus). (credit “mitosis drawings”: modification of work
by Mariana Ruiz Villareal; credit “micrographs”: modification of work by Roy van Heesbeen;
credit “cytokinesis micrograph”: Wadsworth Center/New York State Department of Health;
scale-bar data from Matt Russell)

During prophase, the “first phase,” the nuclear envelope starts to dissociate
into small vesicles, and the membranous organelles (such as the Golgi
complex or Golgi apparatus, and endoplasmic reticulum), fragment and
disperse toward the periphery of the cell. The nucleolus disappears
(disperses). The centrosomes begin to move to opposite poles of the cell.
Microtubules that will form the mitotic spindle extend between the
centrosomes, pushing them farther apart as the microtubule fibers lengthen.
The sister chromatids begin to coil more tightly with the aid of condensin
proteins and become visible under a light microscope.
Reading 4.2 Cell cycle

During prometaphase, the “first change phase,” many processes that were
begun in prophase continue to advance. The remnants of the nuclear
envelope fragment. The mitotic spindle continues to develop as more
microtubules assemble and stretch across the length of the former nuclear
area. Chromosomes become more condensed and discrete. Each sister
chromatid develops a protein structure called a kinetochore in the
centromere region (Figure 8). The proteins of the kinetochore attract and
bind mitotic spindle microtubules. As the spindle microtubules extend from
the centrosomes, some of these microtubules come into contact with and
firmly bind to the kinetochores. Once a mitotic fiber attaches to a
chromosome, the chromosome will be oriented until the kinetochores of
sister chromatids face the opposite poles. Eventually, all the sister
chromatids will be attached via their kinetochores to microtubules from
opposing poles. Spindle microtubules that do not engage the chromosomes
are called polar microtubules. These microtubules overlap each other
midway between the two poles and contribute to cell elongation. Astral
microtubules are located near the poles, aid in spindle orientation, and are
required for the regulation of mitosis.

Figure 8 During prometaphase, mitotic spindle microtubules from opposite poles attach to
each sister chromatid at the kinetochore. In anaphase, the connection between the sister
chromatids breaks down, and the microtubules pull the chromosomes toward opposite
poles.

During metaphase, the “change phase,” all the chromosomes are aligned in
a plane called the metaphase plate, or the equatorial plane, midway
between the two poles of the cell. The sister chromatids are still tightly
attached to each other by cohesin proteins. At this time, the chromosomes
are maximally condensed.
Reading 4.2 Cell cycle

During anaphase, the “upward phase,” the cohesin proteins degrade, and
the sister chromatids separate at the centromere. Each chromatid, now
called a chromosome, is pulled rapidly toward the centrosome to which its
microtubule is attached. The cell becomes visibly elongated (oval shaped) as
the polar microtubules slide against each other at the metaphase plate
where they overlap.

During telophase, the “distance phase,” the chromosomes reach the

opposite poles and begin to decondense (unravel), relaxing into a chromatin
configuration. The mitotic spindles are depolymerized into tubulin monomers
that will be used to assemble cytoskeletal components for each daughter
cell. Nuclear envelopes form around the chromosomes, and nucleosomes
appear within the nuclear area.

Cytokinesis, or “cell motion,” is the second main stage of the mitotic phase,
during which cell division is completed via the physical separation of the
cytoplasmic components into two daughter cells. Division is not complete
until the cell components have been apportioned and completely separated
into the two daughter cells. Although the stages of mitosis are similar for
most eukaryotes, the process of cytokinesis is quite different for eukaryotes
that have cell walls, such as plant cells.

In cells such as animal cells that lack cell walls, cytokinesis starts during late
anaphase. A contractile ring composed of actin filaments forms just inside
the plasma membrane at the former metaphase plate. The actin filaments
pull the equator of the cell inward, forming a fissure. This fissure, or “crack,”
is called the cleavage furrow. The furrow deepens as the actin ring
contracts, and eventually the membrane is cleaved in two (Figure 9).

In plant cells, a new cell wall must form between the daughter cells. During
interphase, the Golgi apparatus accumulates enzymes, structural proteins,
and glucose molecules prior to breaking into vesicles and dispersing
throughout the dividing cell. During telophase, these Golgi vesicles are
transported on microtubules to form a vesicular structure at the metaphase
plate. There, the vesicles fuse and coalesce from the center toward the cell
walls; this structure is called a cell plate. As more vesicles fuse, the cell
plate enlarges until it merges with the cell walls at the periphery of the cell.
Enzymes use the glucose that has accumulated between the membrane
layers to build a new cell wall. The Golgi membranes become parts of the
plasma membrane on either side of the new cell wall (Figure 9).
Reading 4.2 Cell cycle

Figure 9 During cytokinesis in animal cells, a ring of actin filaments forms at the metaphase
plate. The ring contracts, forming a cleavage furrow, which divides the cell in two. In plant
cells, Golgi vesicles coalesce at the former metaphase plate, forming a phragmoplast. A cell
plate formed by the fusion of the vesicles of the phragmoplast grows from the center toward
the cell walls, and the membranes of the vesicles fuse to form a plasma membrane that
divides the cell in two.

Not all cells adhere to the classic cell cycle pattern in which a newly formed
daughter cell immediately enters the preparatory phases of interphase,
closely followed by the mitotic phase. Cells in the G0 phase are not actively
preparing to divide. The cell is in a quiescent (inactive) stage that occurs
when cells exit the cell cycle. Some cells enter G 0 temporarily until an
external signal triggers the onset of G1. Other cells that never or rarely
divide, such as mature cardiac muscle and nerve cells, remain in G 0
permanently.

Prokaryotic cell division

Prokaryotes, such as bacteria, propagate by binary fission. For unicellular
organisms, cell division is the only method to produce new individuals. In
both prokaryotic and eukaryotic cells, the outcome of cell reproduction is a
pair of daughter cells that are genetically identical to the parent cell. In
unicellular organisms, daughter cells are individuals.

Due to the relative simplicity of the prokaryotes, the cell division process,
called binary fission, is a less complicated and much more rapid process than
cell division in eukaryotes. Recall that the single, circular DNA chromosome
of bacteria is not enclosed in a nucleus, but instead occupies a specific
Reading 4.2 Cell cycle

location, the nucleoid, within the cell, and that a prokaryotic cell lacks
organelles but does contain ribosomes, and may contain additional DNA in
the form of small plasmids. The process begins with the replication of the
single, circular chromosome, resulting in two identical copies. As the cell
elongates, the replicated chromosomes move toward opposite ends of the
cell (Figure 10). A new cell wall then forms between the two chromosomes,
dividing the cell into two daughter cells. Each daughter cell receives one
copy of the chromosome and approximately equal amounts of cytoplasm,
ensuring that the genetic material is faithfully passed on to the next
generation.

Figure 10 Binary fission in a prokaryotic cell. 1. The bacterium before binary fission is when
the DNA is tightly coiled. 2. The DNA of the bacterium has replicated. 3. The DNA is pulled to
the separate poles of the bacterium as it increases size to prepare for splitting. 4. The
growth of a new cell wall begins to separate the bacterium. 5. The new cell wall fully
develops, resulting in the complete split of the bacterium. 6. The new daughter cells have
tightly coiled DNA, ribosomes, and plasmids. Credit: Ecoddington14, CC BY-SA 3.0, via
Wikimedia Commons