Professional Documents
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AUTHORS
Douglas Wilkin, Ph.D.
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Niamh Gray-Wilson
interactive content, visit www.ck12.org
CONTRIBUTOR
Jean Brainard, Ph.D.
The names “CK-12” and “CK12” and associated logos and the
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Contents www.ck12.org
Contents
1 Cell 1
1.1 The Cell Theory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.2 Cells - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.3 Discovery of Cells - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.4 Cell Size and Shape - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
1.5 Common Parts of Cells - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
1.6 Cell Structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
1.7 Two Types of Cells - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
1.8 The Nucleus - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
1.9 The Mitochondria - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
1.10 Ribosomes - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
1.11 Endoplasmic Reticulum - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
1.12 The Golgi Apparatus - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
1.13 Vesicles and Vacuoles - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
1.14 Other Structures of Cells - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
1.15 Plant Cells - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
1.16 The Cytoplasm and Cytoskeleton - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
1.17 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
2 Cell Cycle 58
2.1 Cell Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
2.2 Mitosis - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
2.3 Genetic Variation - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
2.4 Meiosis - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
2.5 Significance of Mitosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
2.6 Significance of Meiosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
2.7 Genetic Disorders - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
2.8 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
3 Transport Mechanisms 90
3.1 The Cell Membrane: A Semi-Permeable Barrier . . . . . . . . . . . . . . . . . . . . . . . . . . 91
3.2 Cell Transport - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
3.3 Diffusion - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
3.4 Facilitated Diffusion - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
3.5 Osmosis - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
3.6 Active Transport - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
3.7 Exocytosis and Endocytosis - Advanced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
3.8 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
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C HAPTER
2 Cell Cycle
Chapter Outline
2.1 C ELL C YCLE
2.2 M ITOSIS - A DVANCED
2.3 G ENETIC VARIATION - A DVANCED
2.4 M EIOSIS - A DVANCED
2.5 S IGNIFICANCE OF M ITOSIS
2.6 S IGNIFICANCE OF M EIOSIS
2.7 G ENETIC D ISORDERS - A DVANCED
2.8 R EFERENCES
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Cell division is just one of several stages that a cell goes through during its lifetime. The cell cycle is a repeating
series of events that include growth, DNA synthesis, and cell division. The cell cycle in prokaryotes is quite simple:
the cell grows, its DNA replicates, and the cell divides. In eukaryotes, the cell cycle is more complicated.
The diagram in Figure 2.1 represents the cell cycle of a eukaryotic cell. As you can see, the eukaryotic cell cycle
has several phases. The mitotic phase (M) actually includes both mitosis and cytokinesis. This is when the nucleus
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and then the cytoplasm divide. The other three phases (G1, S, and G2) are generally grouped together as interphase.
During interphase, the cell grows, performs routine life processes, and prepares to divide. These phases are discussed
below. You can watch a eukaryotic cell going through these phases of the cell cycle at the following link: http://w
ww.cellsalive.com/cell_cycle.htm .
FIGURE 2.1
Eukaryotic Cell Cycle. This diagram rep-
resents the cell cycle in eukaryotes. The
First Gap, Synthesis, and Second Gap
phases make up interphase (I). The M
(mitotic) phase includes mitosis and cy-
tokinesis. After the M phase, two cells
result.
Interphase
Interphase of the eukaryotic cell cycle can be subdivided into the following three phases, which are represented in
Figure 2.1:
• Growth Phase 1 (G1): during this phase, the cell grows rapidly, while performing routine metabolic pro-
cesses. It also makes proteins needed for DNA replication and copies some of its organelles in preparation for
cell division. A cell typically spends most of its life in this phase. This phase is sometimes referred to as Gap
1.
• Synthesis Phase (S): during this phase, the cell’s DNA is copied in the process of DNA replication.
• Growth Phase 2 (G2): during this phase, the cell makes final preparations to divide. For example, it makes
additional proteins and organelles. This phase is sometimes referred to as Gap 2.
If the cell cycle occurred without regulation, cells might go from one phase to the next before they were ready.
What controls the cell cycle? How does the cell know when to grow, synthesize DNA, and divide? The cell cycle is
controlled mainly by regulatory proteins. These proteins control the cycle by signaling the cell to either start or delay
the next phase of the cycle. They ensure that the cell completes the previous phase before moving on. Regulatory
proteins control the cell cycle at key checkpoints, which are shown in Figure 2.2. There are a number of main
checkpoints.
• The G1 checkpoint, just before entry into S phase, makes the key decision of whether the cell should divide.
• The S checkpoint determines if the DNA has been replicated properly.
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• The mitotic spindle checkpoint occurs at the point in metaphase where all the chromosomes should have
aligned at the mitotic plate.
FIGURE 2.2
Checkpoints in the eukaryotic cell cycle
ensure that the cell is ready to proceed
before it moves on to the next phase of
the cycle.
Cancer is a disease that occurs when the cell cycle is no longer regulated. This may happen because a cell’s DNA
becomes damaged. Damage can occur due to exposure to hazards such as radiation or toxic chemicals. Cancerous
cells generally divide much faster than normal cells. They may form a mass of abnormal cells called a tumor (see
Figure 2.16). The rapidly dividing cells take up nutrients and space that normal cells need. This can damage tissues
and organs and eventually lead to death.
FIGURE 2.3
These cells are cancer cells, growing out
of control and forming a tumor.
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MEDIA
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URL: https://www.ck12.org/flx/render/embeddedobject/181097
Summary
• The cell cycle is a repeating series of events that cells go through. It includes growth, DNA synthesis, and cell
division. In eukaryotic cells, there are two growth phases, and cell division includes mitosis.
• The cell cycle is controlled by regulatory proteins at three key checkpoints in the cycle. The proteins signal
the cell to either start or delay the next phase of the cycle.
• Cancer is a disease that occurs when the cell cycle is no longer regulated. Cancer cells grow rapidly and may
form a mass of abnormal cells called a tumor.
• See the Cell Cycle at http://www.cellsalive.com/cell_cycle.htm for a detailed summary.
Explore More
Review
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Mitosis, of course. As you can see, mitosis is a multistage process that ensures separation of sister chromatids, and
ensures that daughter cells are just like the parent cell.
Mitosis
Mitosis is the division of the cell’s nucleus, the final step before two daughter cells are produced. Mitosis begins
immediately at the conclusion of interphase, specifically at the end of the G2 phase. The cell enters mitosis as
it approaches its size limitations. Four distinct phases of mitosis have been recognized: prophase, metaphase,
anaphase, and telophase, with each phase merging into the next one (Figure 2.4).
The Phases
Prophase
Prophase is the first and longest phase of mitosis, see Figure 2.5. During prophase, the chromatin (DNA) coils
up into visible chromosomes, each made up of two sister chromatids held together by the centromere. Also during
this phase, the nucleolus disappears, and the spindle begins to form from the centrioles. Most eukaryotic cells
contain structures known as centrosomes, consisting of a pair of centrioles. During prophase, the centrioles begin
to move to opposite ends, or poles, of the cell. As the centrioles migrate, the fiber-like spindle begins to elongate
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2.2. Mitosis - Advanced www.ck12.org
FIGURE 2.4
Mitosis is the phase of the eukaryotic cell
cycle that occurs between DNA replica-
tion and the formation of two daughter
cells. What happens during mitosis? Dur-
ing mitosis, the nucleus divides, paving
the way for two cells to be produced
after cell division, each with a complete
makeup of genetic material.
between the centrioles. The spindle is a thin, cage-like structure made out of microtubules. In plant cells, the spindle
forms without centrioles. The spindle plays an essential role moving chromosomes and in the separation of sister
chromatids.
FIGURE 2.5
The spindle starts to form during
prophase of mitosis. Kinetochores on
the spindle attach to the centromeres of
sister chromatids.
Preprophase
As plant cells have some structural differences compared to an animal cell, an additional stage prior to prophase is
necessary. In plant cells only, prophase is preceded by a preprophase stage. Plant cells have a large central vacuole
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encompassing the center of the cell. Prior to the division of the nucleus, the nucleus must migrate to the center of the
plant cell. To accomplish this, the cell forms a phragmosome, a sheet of cytoplasm that bisects the middle of the
cell. The phragmosome suspends the cell nucleus in the center of the cell in preparation for prophase. Additionally,
during this phase the plane of cell division is established. The accurate control of division planes, which establishes
the placement of the future cell wall, is crucial for the correct structure of plant tissues and organs.
The mitotic spindle also starts to form during preprophase in plant cells. Unlike animal cells, plant cells do not have
centrosomes to organize their mitotic spindles. Instead, in these cells, the nuclear envelope acts as a microtubule
organizing center (MTOC) for spindle formation. The preprophase spindle forms by self-assembly of microtubules
in the cytoplasm surrounding the nuclear envelope.
Prometaphase
During early prometaphase, the nuclear membrane disintegrates and microtubule spindles invade the center of the
cell. Also during this phase, the spindle attaches to the centromere of each chromatid. Specifically, the spindle
attaches to the kinetochore, a protein structure on the centromere where the spindle fibers attach.
Metaphase
During metaphase, the centromeres of the chromosomes line up along the metaphase plate or equatorial plane, in
essence the approximate middle of the cell. This orientation of the chromosomes at the equator of the cell helps to
ensure proper chromosome separation. This alignment allows the spindle fibers to correctly pull the chromatids to
either pole of the cell, resulting in separation of sister chromatids from a chromosome, see Figure 2.6.
FIGURE 2.6
Chromosomes, consisting of sister chromatids, line up at the equator
(metaphase plate) of the cell during metaphase.
Anaphase
Anaphase is the phase in which the sister chromatids separate. The sister chromatids are pulled apart by the
shortening of the microtubules of the spindles, similar to the reeling in of a fish by the shortening of the fishing
line. One sister chromatid moves to one pole of the cell, and the other sister chromatid moves to the opposite
pole. This process occurs when the proteins that bind sister chromatids together are cleaved, resulting in unattached
identical chromosomes, essentially separate daughter chromosomes. These separate chromosomes are pulled apart
by shortening spindle fibers, and pulled toward the centrosomes to which they are attached. At the end of anaphase
the spindle fibers degrade. At this time, each pole of the cell has a complete set of chromosomes, identical to the
amount of DNA at the beginning of G1 of the cell cycle.
Telophase
Telophase is essentially the opposite of prophase and prometaphase. The chromosomes begin to unwind back
into chromatin in preparation to direct the cell’s metabolic activities. A new nucleus forms around each set of
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2.2. Mitosis - Advanced www.ck12.org
chromosomes. This is followed by cytokinesis, the division of the cytoplasm, resulting in two genetically identical
cells, ready to enter G1 of the next cell cycle. The phases of mitosis are summarized in Figure 2.7.
FIGURE 2.7
Mitosis in the Eukaryotic Cell Cycle. Mito-
sis is the multi-phase process in which the
nucleus of a eukaryotic cell divides. In this
diagram, prometaphase is not included as
a separate phase, but incorporated into
prophase.
FIGURE 2.8
This is a representation of dividing plant
cells. Cell division in plant cells differs
slightly from animal cells as a cell wall
must form. Note that most of the cells are
in interphase. Can you find examples of
the different stages of mitosis?
Cytokinesis
Cytokinesis is the final step in cell division. It often occurs concurrently with telophase, though it is a separate
process. Cytokinesis (Figure 2.9) differs between plant and animal cells. In animal cells, the plasma membrane
pinches inward along the cell’s equator until two cells are formed. Specifically, a cleavage furrow containing a
contractile ring develops in approximately the middle of the cell (similar to the position of the metaphase plate),
essentially pinching off the two nuclei and forming separate cells. In plant cells, a cell plate forms along the cells
equator. A new membrane grows along each side of the cell plate, with a new cell wall forming on the outside of
each new membrane.
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At the end of cytokinesis, each daughter cell has a complete copy of the genome of its parent cell. The end of
cytokinesis marks the end of the M-phase, the end of one cell cycle, and the beginning of G1 and interphase of the
next cell cycle.
FIGURE 2.9
Cytokinesis is the final stage of eukaryotic cell division. It occurs differently
in animal (left) and plant (right) cells.
FIGURE 2.10
In this electron micrograph of a cell, the
formation of two new cells is almost com-
plete, as new membrane grows and di-
vides the parent cell.
MEDIA
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Summary
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Review
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Genetic variation. It is this variation that is the essence of evolution. Without genetic differences among individuals,
"survival of the fittest" would not be likely. Everyone would be exactly the same. How would it be determined who
could or would survive? Either all survive, or all perish.
Sexual reproduction results in infinite possibilities of genetic variation. In other words, sexual reproduction results
in offspring that are genetically unique. They differ from both parents and also from each other. This occurs through
a number of mechanisms, including crossing-over, the independent assortment of chromosomes during anaphase I,
and random fertilization.
• When homologous chromosomes form pairs during prophase I of meiosis I, crossing-over can occur. Crossing-
over is the exchange of genetic material between non-sister chromatids of homologous chromosomes. It
results in new combinations of genes on each chromosome.
• When cells divide during meiosis, homologous chromosomes are randomly distributed during anaphase I,
separating and segregating independently of each other. This is called independent assortment. It results in
gametes that have unique combinations of chromosomes.
• In sexual reproduction, two gametes unite to produce an offspring. But which two of the millions of possible
gametes will it be? This is likely to be a matter of chance. It is obviously another source of genetic variation
in offspring. This is known as random fertilization.
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2.3. Genetic Variation - Advanced www.ck12.org
All of these mechanisms working together result in an amazing amount of potential variation. Each human couple,
for example, has the potential to produce more than 64 trillion genetically unique children. No wonder we are all
different!
Crossing-Over
As mentioned above, crossing-over occurs during prophase I, and it is the exchange of genetic material between
non-sister chromatids of homologous chromosomes. Recall during prophase I, homologous chromosomes line up
in pairs, gene-for-gene down their entire length, forming a configuration with four chromatids, known as a tetrad.
The process of pairing the homologous chromosomes is called synapsis. During synapsis, non-sister chromatids
may cross-over at points called chiasmata. Within a chiasma, the genetic material from two non-sister chromatids
actually intertwine around each other, and some material from non-sister chromatids switch chromosomes, that is,
the material breaks off and reattaches at the same position on the homologous chromosome (Figure 2.11). This
exchange of genetic material can happen many times within the same pair of homologous chromosomes, creating
unique combinations of alleles. This process is also known as homologous recombination.
FIGURE 2.11
During crossing-over, segments of DNA
are exchanged between non-sister chro-
matids of homologous chromosomes.
Notice how this can result in an allele (A)
on one chromosome being moved to the
other chromosome. The four chromatids
compose the tetrad, with a chiasma at the
point of exchange.
MEDIA
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In humans, there are over 8 million configurations in which the chromosomes can line up during metaphase I of
meiosis. It is the specific processes of meiosis, resulting in four unique haploid cells, that result in these many
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combinations. This independent assortment, in which the chromosome inherited from either the father or mother
can sort into any gamete, produces the potential for tremendous genetic variation. This process underlies the
chromosomal basis of inheritance. Gregor Mendel’s findings and laws will be discussed in the Inheritance concepts,
but essentially, his findings led to the development of two laws of inheritance: the Law of Segregation and the Law
of Independent Assortment. The Law of Segregation states that when any individual produces gametes, the copies
of a gene separate so that each gamete receives only one copy (one allele) of that gene. The Law of Independent
Assortment states that separate genes for separate traits are passed independently of one another from parents to
offspring.
Together with random fertilization, more possibilities for genetic variation exist between any two people than the
number of individuals alive today. Sexual reproduction is the random fertilization of a gamete from the female
using a gamete from the male. In humans, over 8 million (223 ) chromosome combinations exist in the production
of gametes in both the male and female. Essentially, when the homologous pairs of chromosomes line up during
metaphase I and then are separated at anaphase I, there are (223 ) possible combinations of maternal and paternal
chromosomes. During random fertilization, a sperm cell, with over 8 million possible chromosome combinations,
fertilizes an egg cell, which also has over 8 million possible chromosome combinations. Together, there are over 64
trillion unique combinations, not counting the additional variation produced by crossing-over during prophase I. In
other words, each human couple could produce a child with over 64 trillion unique chromosome combinations!
MEDIA
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Mitosis and meiosis are two types of cell division, with dramatically different products. Mitosis begins with a diploid
somatic cell and ends with two genetically identical diploid cells. Meiosis begins with a diploid cell and produces
four haploid genetically unique cells that form gametes.
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FIGURE 2.12
Mitosis vs. Meiosis Comparison. Mitosis
produces two diploid daughter cells, ge-
netically identical to the parent cell. Meio-
sis produces four haploid daughter cells,
each genetically unique.
MEDIA
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Summary
• Crossing-over, the independent assortment of chromosomes during anaphase I, and random fertilization all
increase the genetic variation of a species.
Review
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