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83

Cell Division
Chapter Concepts
5.1 Maintaining the Chromosome Number
Each species has a characteristic number of
chromosomes. 84
Mitosis (a type of nuclear division) maintains the
chromosome number of cells. Mitosis is
necessary to the growth and repair of body
cells. 84
Mitosis is a part of the cell cycle. First, cells get
ready to divide, and then they divide. 86
5.2 Mitosis in Detail
During mitosis, a complete set of chromosomes
is distributed to each of two daughter cells. 88
5.3 Reducing the Chromosome Number
Meiosis (another type of nuclear division)
reduces the chromosome number in life cycles
involving sexual reproduction. 92
The process of meiosis ensures genetic
recombination in the daughter cells. 92
5.4 Meiosis in Detail
During meiosis, half the total number of
chromosomes is distributed to each of four
daughter cells. 94
5.5 Comparison of Meiosis with Mitosis
Meiosis differs from mitosis both in occurrence
and in process. 96
5.6 The Human Life Cycle
The human life cycle includes both mitosis and
meiosis. 98
In humans, and many other animals, meiosis is a
part of the production of sperm in males and
eggs in females. 98
When the sperm fertilizes the egg, the full
number of chromosomes is restored in
offspring. 98
During mitotic cell division, the daughter cells receive a full
complement of chromosomes. The process is very orderly; it has
to be to ensure that the daughter cells receive one of each
chromosome and not two of one kind and none of another. Cell
division will produce new cells so that the organism can grow.
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skeletal muscle and nervous tissue, do not divide and there-
fore cannot be renewed. Regardless, cell division is con-
trolled and does not usually occur helter-skelter, the notable
exception occurs when cancer, a cell division disease, is
present. Before mitosis takes place, DNA replicates; there-
after, each chromosome is duplicated and has two identical
parts called sister chromatids. Sister chromatids are con-
stricted and attached to each other at a region called the cen-
tromere. During nuclear division the centromere divides
and the sister chromatids separate. Once separation occurs,
they are called daughter chromosomes. These chromo-
somes, which consist of only one chromatid, are distributed
equally to the daughter nuclei. In this way, each daughter
cell gets a copy of each chromosome.
5.1 Maintaining the Chromosome
Number
When a eukaryotic cell is not undergoing division, the DNA
(and associated proteins) within a nucleus is a tangled mass
of thin threads called chromatin. At the time of cell division,
chromatin coils, loops, and condenses to give highly com-
pacted structures that are called chromosomes. The Science
Focus on the next page describes the transition from chro-
matin to chromosomes in greater detail.
When the chromosomes are highly coiled and con-
densed at the time of cell division, it is possible to photo-
graph and count them. Each species has a characteristic
chromosome number; for instance, human cells contain 46
chromosomes, corn has 20 chromosomes, and a craysh has
200! This is called the full or diploid (2n) number of chro-
mosomes that occurs in all cells of the body. The diploid
number includes two chromosomes of each kind. Half the
diploid number is called the haploid (n) number of chromo-
somes, representing only one of each kind of chromosome.
In the life cycle of many animals, only sperm and eggs have
the haploid number of chromosomes.
Cell division in most eukaryotes involves nuclear divi-
sion and cytokinesis, which is division of the cytoplasm.
Somatic, or body, cells undergo mitosisthat is, nuclear
division in which the chromosome number stays constant
(Fig. 5.1). In diploid organisms such as ourselves, a diploid
nucleus divides to produce daughter nuclei that are also
diploid. Some organisms are haploid as adults. In that case,
the haploid nucleus divides to produce daughter nuclei that
are also haploid.
Mitosis is the type of nuclear division that occurs in
growth and repair of the body. Humans begin life as a single
cell but they eventually have one hundred trillion cells as
adults due to mitosis. Even then, mitosis does not stop. As
adults, however, certain tissues such as the epidermis of the
skin, the lining of the digestive and respiratory tracts, and
the lymphoid tissue that produces blood cells account for
much of the cell division that occurs. Other tissues, namely
84 Part 1 Cell Biology 5-2
T
en million times a day a beautiful dance nishes
inside Lauras body. Its the intricate movement of
chromosomes that occurs whenever one of her cells
divides into two. The cell has already duplicated its chromo-
somes. Then, in a carefully choreographed set of maneu-
vers, the two parts of the chromosome separate so that the
resulting two cells each have a complete chromosomal
complement. Through this amazing reproductive process,
skin cells can proliferate to repair a wound and the immune
system can quickly amass an army of cells to defeat an
infection. And through a renement of the basic cell division
process, sperm and egg cells arise, ready to join and create
a new life. This chapter will provide a guided tour of cells as
they divide.
DNA replication
Mitosis
during interphase
centromere
2n = 4
2n = 4
sister
chromatids
2n = 4 2n = 4
chromosome
centriole
Figure 5.1 Mitosis overview.
Following DNA replication during interphase, each chromosome in
the parental nucleus is duplicated and consists of two sister
chromatids. During mitosis, the centromeres divide and the sister
chromatids separate, becoming daughter chromosomes that move
into the daughter nuclei. Therefore, daughter cells have the same
number and kinds of chromosomes as the parental cell. (The blue
chromosomes were inherited from the father, and the red
chromosomes were inherited from the mother.)
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When early investigators decided that the genes are contained
in the chromosomes, they had no idea of chromosome composi-
tion. By the mid-1900s, it was known that chromosomes are
made up of both DNA and protein. Only in recent years, how-
ever, have investigators been able to produce models suggesting
how chromosomes are organized.
A eukaryotic chromosome is more than 50% protein. Many
of these proteins are concerned with DNA and RNA synthesis,
but a large proportion, termed histones, seem to play primarily
a structural role. There are ve primary types of histone mole-
cules, designated H1, H2A, H2B, H3, and H4. Remarkably, the
amino acid sequences of H3 and H4 vary little between organ-
isms. For example, the H4 of peas is only two amino acids dif-
ferent from the H4 of cattle. This similarity suggests that there
have been few mutations in the histone proteins during the
course of evolution and that the histones therefore have very
important functions.
Ahuman cell contains at least 2 meters of DNA. Yet all of this
DNA is packed into a nucleus that is about 5 m in diameter.
The histones are responsible for packaging the DNA so that it
can t into such a small space. First the DNA double helix is
wound at intervals around a core of eight histone molecules
(two copies each of H2A, H2B, H3, and H4), giving the appear-
ance of a string of beads (Fig. 5Aa and b). Each bead is called a
nucleosome, and the nucleosomes are said to be joined by
linker DNA. This string is coiled tightly into a ber that has
six nucleosomes per turn (Fig. 5Ac). The H1 histone appears to
mediate this coiling process. The ber loops back and forth (Fig.
5Ad and e) and can condense to produce a highly compacted
form (Fig. 5Af ) characteristic of metaphase chromosomes. No
doubt, compact chromosomes are easier to move about than
extended chromatin.
85
Whats in a Chromosome?
Figure 5A Levels of chromosome structure.
Each drawing has a scale giving a measurement of length for that
drawing. Notice that each measurement represents an ever-
increasing length; therefore, it would take a much higher
magnication to see the structure in (a) than in (f).
Figure 5B Eukaryotic nucleus.
The nucleus contains chromatin, DNA at two different levels of
coiling and condensation. Euchromatin is at the level of looped
chromatin, and heterochromatin is at the level of condensed
chromatin in Figure 5A. Arrows indicate nuclear pores.
c.
d.
e.
f.
euchromatin
heterochromatin
3
0

n
m
3
0
0

n
m
7
0
0

n
m
nucleosome
nucleosome
histones
a. DNA helix
b. Nucleosomes
Coiled
nucleosomes
Looped
chromatin
Condensed
chromatin
Condensed
chromosome
2

n
m
1
1

n
m
1
,
4
0
0

n
m
histone
H1
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Cells differ in the length of time it takes them to com-
plete the cell cycle. The difference seems to depend on how
long they spend in G
1
. There are even some human cells
such as nerve cells and skeletal muscle cells that become
permanently arrested in G
1
, and these cells are said to have
entered a G
0
stage. Once a mammalian cell enters the S
stage, it usually only takes about 12 to 24 hours to nish the
cell cycle.
The term interphase is now used to mean all the stages
of the cell cycle (i.e., stages G
1
, S, and G
2
) except mitosis and
cytokinesis. One thing to keep clearly in mind is that DNA
replication occurs during the S stagethat is during inter-
phase.
Cells undergo a cycle that includes the G
1
, S, G
2
,
and M stages.
Proteins Regulate the Cell Cycle
If a cell arrested in the G
0
stage is placed in the cytoplasm of
an S-stage cell, it will go on and nish the cell cycle. Similar-
ly, there are cells that enter and never get beyond the G
2
stage. If this type of cell is fused with a cell undergoing mito-
sis, it will go ahead and undergo mitosis. From this data,
researchers deduced that there are two places in the cell
cycle where stimulatory proteins are needed to make the cell
nish the complete cell cycle:
G
1
stage S stage when DNAis synthesized
G
2
stage M stage when mitosis occurs
Over the past few years, researchers have made
remarkable progress identifying the proteins that cause a
cell to move from the G
1
stage to the S stage and the pro-
teins that cause a cell to move from the G
2
stage to the M
stage. Some of these biologists worked with frog eggs, oth-
ers with yeast cells, and still others used cell cultures as
their experimental material. The researchers identied two
types of proteins of interest: kinases and cyclins. Akinase is
an enzyme that removes a phosphate group from ATP and
adds it to another protein. The addition of the phosphate
group to that protein activates it. Activation by a kinase is a
common way for cells to turn on a cellular process, but it
turned out that the kinases involved in the cell cycle are
themselves activated when they combine with a protein
called cyclin. Cyclins are so named because their quantity is
not constant in the cell.
Figure 5.3 is an illustration that shows the process in a
clockwise diagram. After S-kinase combines with S-cyclin,
the kinase phosphorylates a protein that causes the cell to
move from the G
1
stage to the S stage when DNAis synthe-
The Cell Cycle
Because there was little visible activity between divisions,
early investigators dismissed this period of time as a rest-
ing state termed interphase. When it was discovered in
the 1950s that DNA replication and chromosome duplica-
tion occur during interphase, the cell cycle concept was
proposed.
Cells grow and divide during a cycle that has four
stages (Fig. 5.2). The entire cell division stage, including
both mitosis and cytokinesis, is termed the M stage
(M mitosis). The period of DNA synthesis when replica-
tion occurs is termed the S stage (S synthesis) of the cycle.
There are two other stages of the cycle. The period of
time prior to the S stage is termed the G
1
stage, and the peri-
od of time prior to the M stage is termed the G
2
stage. During
the G
1
stage, the cell grows in size and the cellular organelles
increase in number. During the G
2
stage, various metabolic
events occur in preparation for mitosis. When rst designat-
ed, G meant gap, but some biologists now prefer
G growth.
86 Part 1 Cell Biology 5-4
I
n
t
e
r
p
h
a
s
e
G
1
S
G
2
organelles begin
to double
in number
replication
of DNA
synthesis of
proteins
Time
M
i
t
o
s
i
s
c
y
to
k
in
e
s
is
telophase
anaphase
metaphase
p
ro
p
h
a
s
e
Figure 5.2 The cell cycle.
Cells go through a cycle that consists of four stages. The length of
time that cells take to nish the cell cycle varies and some
specialized cells like nerve cells and skeletal muscle cells no longer
progress through the cycle. Mammalian cells usually take at least 16
hours to complete the cell cycle.
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sized (replicated). After that occurs, S-cyclin is destroyed,
and S-kinase is no longer active.
Similarly, after M-kinase combines with M-cyclin, the
kinase phosphorylates a protein that causes the cell to move
from the G
2
stage to the M stage when mitosis occurs. Three
things occur: (1) chromosomes condense, (2) the nuclear
envelope disassembles, and (3) the spindle forms. (The spin-
dle is the structure involved in chromosome movement dur-
ing mitosis.) Now M-cyclin is destroyed.
Until recently, the mechanics of the cell cycle and the
causes of cancer were thought to be distantly related. Now
they appear to be intimately related. Growth factors are mol-
ecules that attach to plasma membrane receptors and there-
by bring about cell growth. Ordinarily, a cyclin might
combine with its kinase only when a growth factor is
Chapter 5 Cell Division 87 5-5
P
mitosis
Time
G
1
S
G
2
M-cyclin
M-cyclin is
destroyed.
S-kinase
S-cyclin
S-cyclin is
destroyed.
M-kinase
S-kinase combines
with S-cyclin.
S-kinase phosphorylates
a protein and the product
symbolized as
triggers DNA
synthesis.
M
M-kinase phosphorylates
a protein and the product
symbolized as
triggers mitosis.
M-kinase combines
with M-cyclin.
P
P
P
present. But a cyclin that has gone awry might combine with
its kinase even when a growth factor is not present. The
result would be a tumor. There are genes called tumor-
suppressor genes that usually function to prevent cancer
from occurring. It has been shown that the product of one
major tumor-suppressor gene (known as p53) brings about
the production of a protein that combines with kinases and
prevents them from becoming activated.
Proteins regulate the cell cycle. A kinase combines
with a cyclin at two critical checkpoints in the cell
cycle: the beginning of the S stage when DNA is
synthesized and the M stage with mitosis begins.
Figure 5.3 Control of the cell cycle.
At two critical checkpoints a kinase combines with a cyclin. Just before the S stage, S-kinase combines with S-cyclin and synthesis (replication)
of DNA takes place. Just before the M stage, M-kinase combines with M-cyclin and mitosis takes place.
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centrosome. The fact that plant cells lack centrioles suggests
that centrioles are not required for spindle formation.
Mitosis in Animal Cells
Mitosis is a continuous process that is arbitrarily divided into
four phases for convenience of description. These phases are
prophase, metaphase, anaphase, and telophase (Fig. 5.5).
Prophase
It is apparent during early prophase that cell division is
about to occur. The centrosomes begin moving away from
each other toward opposite ends of the nucleus. Spindle
bers appear between the separating centrosomes as the
nuclear envelope begins to fragment, and the nucleolus
begins to disappear.
The chromosomes are now visible. Each is duplicated and
composed of sister chromatids held together at a centromere.
88 Part 1 Cell Biology 5-6
5.2 Mitosis in Detail M
Mitosis is nuclear division that produces two daughter nuclei,
each with the same number and kinds of chromosomes as the
parental nucleus.
During mitosis, a spindle brings about an orderly distri-
bution of chromosomes to the daughter cell nuclei. The spin-
dle contains many bers, each composed of a bundle of
microtubules. Microtubules are hollow cylinders found in
the cytoplasm which can assemble and disassemble. When
microtubules assemble, tubulin protein dimers come togeth-
er, and when they disassemble, the tubulin dimers separate.
The centrosome, which is the main microtubule-
organizing center of the cell, divides before mitosis begins
(Fig. 5.4). Its believed that centrosomes are responsible for
organizing the spindle. Each centrosome contains a pair of
barrel-shaped organelles called centrioles and an aster
which is an array of short microtubules that radiate from the
Early Prophase
Chromosomes are duplicated.
Centrosomes begin moving apart;
nuclear envelope is fragmenting
and nucleolus will disappear.
Late Prophase
Spindle is in process of forming, and
centromeres of chromosomes are
attaching to centromeric spindle
fibers.
Figure 5.4 Late interphase. Figure 5.5 Phases of animal cell mitosis.
chromatin
aster
centrioles
in centrosomes
nucleolus
nuclear
envelope
plasma
membrane
Late Interphase
Chromatin is condensing into
chromosomes and centrosomes
have duplicated in preparation
for mitosis.
20 m
centrioles in centrosome
chromosome
centromere
nucleolus
aster
centromeric
spindle fiber
plasma
membrane
chromosomes
centromere
20 m 9 m
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Counting the number of centromeres in diagrammatic draw-
ings tells you the number of chromosomes in a cell.
The spindle begins forming during late prophase and the
chromosomes become attached to the spindle bers. Their
centromeres attach to bers called centromeric (also called
kinetochore) bers. As yet, the chromosomes have no particu-
lar orientation as they move rst one way and then the other.
Metaphase
By the time of metaphase, the fully formed spindle consists
of poles, asters, and bers. The chromosomes attached to
centromeric spindle bers are aligned at the metaphase
plate (also called the equator) of the spindle. Polar spindle
bers reach beyond the metaphase plate and overlap. At the
close of metaphase, the centromeres uniting the sister chro-
matids divide.
Anaphase
During anaphase, the centromeres divide. The sister chro-
matids separate, becoming two daughter chromosomes that
move toward the opposite poles of the spindle. The daugh-
ter chromosomes have a centromere and single chromatid.
What accounts for the movement of the daughter chromo-
somes? First, the centromeric spindle bers disassemble at
the region of the kinetochore, and this pulls the daughter
chromosomes to the poles. Second, the polar spindle bers
lengthen as they slide past one another.
Telophase
During telophase, the spindle disappears and nuclear enve-
lope components reassemble around the daughter chromo-
somes. Each daughter nucleus contains the same number
and kinds of chromosomes as the original parental cell.
Remnants of the polar spindle bers are still visible between
the two nuclei.
The chromosomes become more diffuse chromatin once
again, and a nucleolus appears in each daughter nucleus.
Cytokinesis is nearly complete, and soon there will be two
individual daughter cells, each with a nucleus that contains
the diploid number of chromosomes.
Chapter 5 Cell Division 89 5-7
daughter
chromosome
Daughter chromosomes (each
consisting of one chromatid)
are moving toward the
poles of the spindle.
Anaphase
nucleolus
Daughter cells are forming as nuclear
envelopes and nucleoli appear.
Chromosomes will become indistinct
chromatin.
Telophase
cleavage
furrow
chromosomes
at
metaphase
plate
Chromosomes (each consisting of
two sister chromatids) are at the
metaphase plate (center of fully
formed spindle).
polar spindle
fiber
Metaphase
aster
pole
20 m 16 m 20 m
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How Plant Cells Divide
As with animal cells, mitosis in plant cells permits growth
and repair. Certain plant tissue, called meristematic tissue,
retains the ability to divide throughout the life of a plant.
Meristematic tissue is found in root tip and shoot tip of
stems. Lateral meristem accounts for the ability of trees to
increase their girth each growing season.
Figure 5.6 illustrates mitosis in plant cells; exactly the
same phases are seen in plant cells as in animal cells. During
early prophase, the chromatin condenses into scattered pre-
viously duplicated chromosomes and the spindle forms;
during late prophase, chromosomes attach to spindle bers;
during metaphase, the chromosomes are at the metaphase
plate of the spindle; during anaphase, the sister chromatids
separate becoming daughter chromosomes that move into
the daughter nuclei; and during anaphase, cytokinesis
begins. Although plant cells have a centrosome, and spindle,
there are no centrioles nor asters during cell division.
Mitosis in plant and animal cells ensures the
daughter cells have the same number and kinds of
chromosomes as the parental cell.
90 Part 1 Cell Biology 5-8
Figure 5.6 Phases of plant cell mitosis
Note the absence of centrioles and asters and the presence of the cell wall. In telophase, a cell plate develops between the two daughter cells.
The cell plate marks the boundary of the new daughter cells, where new plasma membrane and a new cell wall is forming for each cell.
Cytokinesis in Plant and Animal Cells
Cytokinesis, or cytoplasmic cleavage, usually accompanies
mitosis. Division of the cytoplasm begins in anaphase, con-
tinues in telophase, but does not reach completion until just
before the following interphase. By that time, the newly
forming cells have received a share of the cytoplasmic
organelles which duplicated during the previous interphase.
Cytokinesis in Plant Cells
Cytokinesis in plant cells occurs by a process different from
that seen in animal cells (Fig. 5.7). The rigid cell wall that
surrounds plant cells does not permit cytokinesis by furrow-
ing. Instead, the Golgi apparatus produces membranous
sacs called vesicles, which move along the microtubules to
the midpoint between the two daughter nuclei. These vesi-
cles fuse, forming a cell plate. Their membrane completes
the plasma membrane for both cells. They also release mole-
cules that signal the formation of plant cell walls, which are
strengthened by the addition of cellulose brils.
A spindle forms during mitosis in plant cells, but
there are no centrioles or asters. Cytokinesis in
plant cells involves the formation of a cell plate.
20 m
20 m
20 m
20 m Prophase
Metaphase
Anaphase
Telophase
cell wall
cell plate
chromosomes
spindle
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Cytokinesis in Animal Cells
In animal cells, a cleavage furrow, which is an indentation of
the membrane between the two daughter nuclei, begins as
anaphase draws to a close. The cleavage furrow deepens
when a band of actin laments, called the contractile ring,
slowly forms a constriction between the two daughter cells.
The action of the contractile ring can be likened to pulling a
drawstring ever tighter about the middle of a balloon. As the
drawstring is pulled tight, the balloon constricts in the middle.
Anarrow bridge between the two cells can be seen dur-
ing telophase, and then the contractile ring continues to sepa-
rate the cytoplasm until there are two daughter cells (Fig. 5.8).
Cytokinesis in animal cells is accomplished by a
furrowing.
Cell Division in Prokaryotes
Asexual reproduction requires a single parent, and the off-
spring are identical to the parent because they contain the
same genes. The process of asexual reproduction in prokary-
otes is termed binary ssion because division (ssion) pro-
duces two (binary) daughter cells that are identical to the
original parental cell. Before division occurs, DNAreplicates
and the single chromosome is duplicated. Thus, there are
two chromosomes that separate as the cell elongates. When
the cell is approximately twice its original length, the plasma
membrane grows inward and a new cell wall forms, divid-
ing the cell into two approximately equal portions.
Asexual reproduction in prokaryotes is by binary
ssion. Following DNA replication, the two resulting
chromosomes separate as the cell elongates.
Chapter 5 Cell Division 91 5-9
vesicles containing
membrane components
fusing to form cell plate
nuclei cell wall
cleavage furrow
contractile ring
Figure 5.7 Cytokinesis in plant cells.
During cytokinesis in a plant cell, the cell plate forms midway
between the two daughter nuclei and extends to the plasma
membrane.
Figure 5.8 Cytokinesis in an animal cell.
A single cell becomes two cells by a furrowing process. A contractile
ring composed of actin laments gradually gets smaller, and the
cleavage furrow pinches the cell into two cells.


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5.3 Reducing the Chromosome
Number
Meiosis occurs in any life cycle that involves sexual repro-
duction. Meiosis reduces the chromosome number in such a
way that the daughter nuclei receive only one member of
each homologous pair. The process of meiosis ensures that
the next generation of individuals will have a combination
of traits that are different from either parent.
Overview of Meiosis
Meiosis requires two nuclear divisions and produces four haploid
daughter cells, each having one of each kind of chromosome and
therefore half the total number of chromosomes present in the
diploid parental nucleus. The parental cell has the diploid
number of chromosomes, while the daughter cells have the
haploid number of chromosomes.
Recall that when a cell is 2n or diploid, the chromo-
somes occur in pairs. For example, the 46 chromosomes of
humans occur in 23 pairs of chromosomes. The members of
each pair are called homologous chromosomes or homo-
logues.
Figure 5.9 presents an overview of meiosis, indicating
the two cell divisions, meiosis I and meiosis II. Prior to
meiosis I, DNA replication has occurred and the chromo-
somes are duplicated. Each chromosome consists of two
chromatids held together at a centromere. During meiosis I
the homologous chromosomes come together and line up
side by side. This so-called synapsis results in an association
of four chromatids that stay in close proximity during the
rst two phases of meiosis I.
Due to synapsis there are pairs of homologous chromo-
somes at the metaphase plate during meiosis I. Then, the
members of these pairs separate and each daughter cell
receives one member of each pair. Therefore, the daughter
cells have the haploid number of chromosomes, as you can
verify by counting the number of centromeres. Each chro-
mosome, however, is still duplicated.
During meiosis I, homologous chromosomes pair
up and then separate. Each daughter cell receives
one copy of each kind of chromosome.
No replication of DNAis needed between meiosis I and
meiosis II because the chromosomes are already duplicated:
they already have two sister chromatids. During meiosis II,
the centromeres divide and the sister chromatids separate,
becoming daughter chromosomes that are distributed to
daughter nuclei. In the end, each of four daughter cells has
the haploid number of chromosomes and each chromosome
consists of one chromatid.
Following meiosis II, there are four haploid
daughter cells and each chromosome consists of
one chromatid.
In some life cycles, such as that of humans (see Fig.
5.15), the daughter cells mature into gametes (sex cells
sperm and egg) that fuse during fertilization. Fertilization
restores the diploid number of chromosomes in a cell that
will develop into a new individual.
92 Part 1 Cell Biology 5-10
DNA
replication
Meiosis I
Meiosis II
centromere
paternal member
of pair nucleolus
maternal member
of pair
synapsis
n = 2
2n = 4
2n = 4
n = 2 n = 2
n = 2
sister
chromatids
homologous pairs
separate
sister chromatids
separate, becoming
daughter chromosomes
Figure 5.9 Overview of meiosis.
Following DNA replication, each chromosome is duplicated. During
meiosis I, the homologous chromosomes pair during synapsis and
then separate. During meiosis II, the centromeres divide and the
sister chromatids separate, becoming daughter chromosomes that
move into the daughter nuclei.
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Chapter 5 Cell Division 93 5-11
orientations are considered, the result will be 2
3
or eight
combinations of maternal and paternal chromosomes in the
resulting gametes from this cell. In humans, where there are
23 pairs of chromosomes, the possible chromosomal combi-
nations in the gametes is a staggering 2
23
, or 8,388,608. And
this does not even consider the genetic variations that are
introduced due to crossing-over.
During meiosis, crossing-over mixes the genetic
information of maternal and paternal
chromosomes and independent assortment leads
to different combinations of these chromosomes in
the gametes and offspring.
Genetic Recombination
Meiosis helps ensure that genetic recombination occurs
through two key events: crossing-over and independent
assortment of homologous chromosomes. In order to
appreciate the signicance of these events, it is necessary to
realize that the members of a homologous pair can carry
slightly different instructions for the same genetic trait. For
example, one homologue may carry instructions for brown
eyes while the corresponding homologue may carry
instructions for blue eyes.
Crossing-over of Nonsister Chromatids
Its often said that we inherit half our chromosomes from
our mother and half from our father, but this is not strictly
correct because of crossing-over. During synapsis, the
homologous chromosomes come together and line up side
by side. Now an exchange of genetic material may occur
between the nonsister chromatids of the homologous pair
(Fig. 5.10). Crossing-over means that the genetic instruc-
tions from a mother and father are mixed and the chro-
matids held together by a centromere are no longer
identical. When the chromatids separate during meiosis I,
the daughter cells receive chromosomes with recombined
genetic material.
Independent Assortment of Homologous Chromosomes
Independent assortment means that the homologous chro-
mosomes separate independently or in a random manner.
When homologues align at the metaphase plate, the mater-
nal or paternal homologue may be orientated toward either
pole. Figure 5.11 shows four possible orientations for a cell
that contains only three pairs of chromosomes. Each orien-
tation results in gametes that have a different combination
of maternal and paternal chromosomes. Once all possible
sister chromatids
synapsis
crossing-over
between nonsister
chromatids
chromatids
after
exchange
recombinant
daughter
chromosomes
Figure 5.10 Synapsis and crossing-over.
During meiosis I, from left to right, duplicated homologous
chromosomes undergo synapsis; nonsister chromatids break and
then rejoin, so that two of the resulting daughter chromosomes have
a different combination of genes.
Figure 5.11 Independent assortment.
Four possible orientations of homologue pairs at the metaphase plate are shown. Each of these will result in daughter nuclei with a different
combination of parental chromosomes. When a cell has three pairs of homologous chromosomes, there are 23 possible combinations of parental
chromosomes in the daughter nuclei.
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94 Part 1 Cell Biology 5-12
5.4 Meiosis in Detail M
The same four phases seen in mitosisprophase,
metaphase, anaphase, and telophaseoccur during both
meiosis I and meiosis II.
The First Division
Phases of meiosis for an animal cell are diagrammed in Fig-
ure 5.12. During prophase I, the spindle appears while the
nuclear envelope fragments and the nucleolus disappears.
Due to DNAduplication during interphase, the homologous
chromosomes each have two sister chromatids. During
synapsis, crossing-over can occur. If so, the sister chromatids
of a duplicated chromosome are no longer identical.
During metaphase I, homologous pairs are aligned at
the metaphase plate. The maternal homologue may be ori-
entated toward either pole, and the father homologue may
be aligned toward either pole. This means that all possible
combinations of chromosomes can occur in the daughter
nuclei. During anaphase I, homologous chromosomes sep-
arate and move to opposite poles of the spindle. Each chro-
mosome still consists of two chromatids.
In some species, there is a telophase I phase at the end of
meiosis I. If so, the nuclear envelopes re-form and nucleoli
appear. This phase may or may not be accompanied by
cytokinesis, which is separation of the cytoplasm.
No replication of DNA occurs during a period of time
between divisions called interkinesis.
The Second Division
Phases of meiosis II for an animal cell are diagrammed in
Figure 5.13. At the beginning of prophase II, a spindle
appears while the nuclear envelope dissembles and the
nucleolus disappears. Each duplicated chromosome is
attached to the spindle and lines up independently at the
metaphase plate during metaphase II. At the close of
metaphase II, the centromeres divide. During anaphase II,
sister chromatids separate, becoming daughter chromo-
somes that move into the daughter nuclei. In telophase II,
the spindle disappears as nuclear envelopes re-form. The
plasma membrane furrows to give two complete cells, each
of which has the haploid number of chromosomes. Each
chromosome consists of one chromatid. Since each cell from
meiosis I undergoes meiosis II, there are four daughter cells
altogether.
During meiosis I, crossing-over occurs.
Homologous chromosomes, each consisting of
two sister chromatids, separate, and the daughter
cells are haploid. Following meiosis II, there are
four haploid daughter cells, and each chromosome
has only one chromatid.
Prophase I
Homologous pairs
during synapsis.
Metaphase I
Homologous pairs
align at the metaphase
plate.
Anaphase I
Homologous chromosomes
separate, pulled to opposite
poles by centromeric
spindle fibers.
Telophase I
Daughter cells have
one chromosome from
each homologous pair.
Interkinesis
Chromosomes still
consist of
two chromatids.
DNA
replication
2n = 4
n = 2
Meiosis I
Figure 5.12 Meiosis I.
The exchange of color between nonsister chromatids represents
crossing-over.
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Overview
Chapter 5 Cell Division 95 5-13
Figure 5.13 Meiosis II.
During meiosis II sister chromatids separate, becoming daughter chromosomes that are distributed to the daughter nuclei. Following meiosis II,
there are four haploid daughter cells. Comparing the number of centromeres in the daughter cells with the number in the parental cell at the start
of meiosis I veries that the daughter cells are haploid.
Meiosis II
Prophase II
Cells have one
chromosome
from each
homologous pair.
Metaphase II
Chromosomes align
at the metaphase
plate.
Anaphase II
Daughter chromosomes
move toward the poles.
Telophase II
Spindle disappears,
nuclei form, and
cytokinesis takes
place.
Daughter Cells
Meiosis results
in four haploid
daughter cells.
n = 2 n = 2
n = 2 n = 2
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5.5 Comparison of Meiosis with
Mitosis
Figure 5.14 compares mitosis to meiosis. The differences
between these cellular divisions can be categorized accord-
ing to occurrence and process.
Occurrence
Meiosis occurs only at certain times in the life cycle of sexu-
ally reproducing organisms. In humans, meiosis occurs only
in the reproductive organs and produces the gametes. Mito-
sis is more common because it occurs in all tissues during
growth and repair.
Process
We will compare both meiosis I and meiosis II to mitosis.
Comparison of Meiosis I to Mitosis
The following are distinctive differences between the
processes of meiosis I and mitosis.
3. Homologous chromosomes (with centromeres intact)
separate and move to opposite poles during anaphase I
in meiosis. Sister chromatids separate, becoming
daughter chromosomes that move to opposite poles
during anaphase in mitosis.
Comparison of Meiosis II to Mitosis
The events of meiosis II are just like those of mitosis except that
in meiosis II, the nuclei contain the haploid number of chro-
mosomes. The following listing compares meiosis II to mitosis:
96 Part 1 Cell Biology 5-14
Meiosis I
Prophase I
Pairing of chromosomes
Metaphase I
Homologous chromosomes
at metaphase plate
Anaphase I
Homologous chromosomes
separate
Telophase I
Daughter cells are haploid
Mitosis
Prophase
No pairing of chromosomes
Metaphase
Duplicated chromosomes at
metaphase plate
Anaphase
Sister chromatids separate,
becoming daughter chromo-
somes that move to the
poles
Telophase
Daughter cells are diploid
Meiosis II
Prophase II
No pairing of chromosomes
Metaphase II
Haploid number of
duplicated chromosomes at
metaphase plate
Anaphase II
Sister chromatids separate,
becoming daughter
chromosomes that move to
the poles
Telophase II
Four daughter cells
Mitosis
Prophase
No pairing of chromosomes
Metaphase
Diploid number of
duplicated chromosomes at
metaphase plate
Anaphase
Sister chromatids separate,
becoming daughter
chromosomes that move to
the poles
Telophase
Two daughter cells
These events distinguish meiosis I from mitosis.
1. Homologous chromosomes pair and undergo crossing-
over during prophase I of meiosis but not during
mitosis.
2. Paired homologous chromosomes align at the
metaphase plate during metaphase I in meiosis.
Individual (duplicated chromosomes) align at the
metaphase plate during metaphase in mitosis.
The following are differences between meiosis and mitosis:
1. DNAreplication takes place only once during both
meiosis and mitosis. There are two nuclear divisions
during meiosis and only one nuclear division during
mitosis.
2. Four daughter cells are produced by meiosis. Mitosis
results in two daughter cells.
3. The four daughter cells formed by meiosis are haploid.
The daughter cells produced by mitosis have the same
chromosome number as the parental cell.
4. The daughter cells from meiosis are not genetically
identical to each other or to the parental cell. The
daughter cells from mitosis are genetically identical to
each other and to the parental cell.
Meiosis is a specialized process that reduces the
chromosome number and occurs only during the
production of gametes. Mitosis is a process that
occurs during growth and repair of all tissues.
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Chapter 5 Cell Division 97 5-15
Meiosis Mitosis
homologues align
independently
synapsis and
crossing-over
occur
homologues separate
sister chromatids
separate
chromosomes align
at the metaphase plate
daughter cells form
sister
chromatids
separate
daughter nuclei are not genetically
identical to parent cell
daughter nuclei are genetically
identical to parent cell
daughter
cells form
Figure 5.14 Meiosis compared to mitosis.
Why does meiosis produce haploid daughter cells while mitosis produces diploid daughter cells? Compare metaphase I of meiosis to metaphase
of mitosis. Only in metaphase I are the homologous chromosomes paired at the metaphase plate. Members of the homologous chromosomes
separate during anaphase I, and therefore the daughter cells are haploid. The blue chromosomes were inherited from one parent and the red
chromosomes were inherited from the other parent. The exchange of color between nonsister chromatids represents crossing-over during
meiosis I.
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Comparison View
5.6 The Human Life Cycle M
The human life cycle requires both meiosis and mitosis (Fig.
5.15). In human males, meiosis is a part of spermatogenesis
which occurs in the testes and produces sperm. In human
females, meiosis is a part of oogenesis which occurs in the
ovaries and produces eggs (Fig. 5.15). Ahaploid sperm and
a haploid egg join at fertilization and the resulting zygote
has the full or diploid number of chromosomes. During
development of the fetus, which is the stage of development
before birth, mitosis keeps the chromosome number con-
stant in all the cells of the body. After birth, mitosis is
involved in the continued growth of the child and repair of
tissues at any time. As a result of mitosis, each somatic cell in
the body has the same number of chromosomes.
Spermatogenesis and Oogenesis in Humans
Spermatogenesis is the production of sperm in males, and
oogenesis is the production of eggs in females. In the testes
of human males, primary spermatocytes, which are diploid
(2n), divide during the rst meiotic division to form two sec-
ondary spermatocytes, which are haploid (n). Secondary
spermatocytes divide during the second meiotic division to
produce four spermatids which are also haploid (n). Whats
the difference between the chromosomes in haploid sec-
ondary spermatocytes and those in haploid spermatids? The
chromosomes in secondary spermatocytes are duplicated
and consist of two chromatids, while those in spermatids
consist of only one chromatid. Spermatids mature into
sperm (spermatozoa). In human males, sperm have 23 chro-
mosomes, which is the haploid number. The process of
meiosis in males always results in four cells that become
sperm.
In ovaries of human females, a primary oocyte, which is
diploid (2n), divides during the rst meiotic division into
two cells, each of which is haploid but the chromosomes are
duplicated. One of these cells, termed the secondary oocyte,
receives almost all the cytoplasm. The other is a polar body.
A polar body is a nonfunctioning cell that occurs during
oogenesis. It contains little cytoplasm and will eventually
disintegrate. The secondary oocyte begins the second meiot-
ic division but stops at metaphase II. The secondary oocyte
leaves the ovary and enters an oviduct where it may be
98 Part 1 Cell Biology 5-16
mitosis
meiosis
meiosis
2n 2n
2n
n
n
fertilization
zygote
2n
mitosis
Figure 5.15 Life cycle of humans.
Meiosis in human males is a part of sperm production, and meiosis in human females is a part of egg production. When a haploid sperm fertilizes
a haploid egg, the zygote is diploid. The zygote undergoes mitosis as it develops into a newborn child. Mitosis continues after birth until the
individual reaches maturity; then the life cycle begins again.
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approached by a sperm. If a sperm does enter the oocyte, the
oocyte is activated to complete the second meiotic division.
The mature egg has 23 chromosomes, each consisting of one
chromatid. In human females, meiosis produces only one
egg and two polar bodies. The polar bodies are a way to dis-
card unnecessary chromosomes while retaining much of the
cytoplasm in the egg. The cytoplasm serves as a source of
nutrients for the developing embryo.
Genetic Recombination in Humans
Notice that fertilization is another means by which chromo-
somes are recombined in the next generation. Because each
child receives both paternal and maternal chromosomes, no
child is exactly like either parent. Altogether there are three
ways in which meiosis ensures that a child has a different
combination of genes than either parent.
1. Independent assortment of chromosomes means that
all possible combinations of chromosomes occur in the
gametes.
2. Crossing-over recombines genetic material so that
sister chromatids are genetically dissimilar.
3. Upon fertilization, recombination of chromosomes
occurs.
Sexual reproduction ensures that each generation
has the same number of chromosomes, and that
each individual has a different genetic makeup
than either parent.
Chapter 5 Cell Division 99 5-17
Spermatogenesis
Oogenesis
primary
spermatocyte
(2n)
primary
oocyte
(2n)
secondary
oocyte
(n)
egg (n) zygote (2n)
first polar
body (n)
meiosis I completion of
meiosis II
meiosis I
meiosis II
secondary
spermatocytes
(n)
spermatids
(n)
sperm
(n)
first polar
body (n)
second polar
body (n)
sperm
nucleus
(n)
fusion of
sperm nucleus (n)
and egg nucleus (n)
nucleus
fertilization
maturation
Figure 5.16 Spermatogenesis and oogenesis.
Spermatogenesis produces four viable sperm, whereas oogenesis produces one egg and two polar bodies. In humans, both sperm and egg have
23 chromosomes each; therefore, following fertilization, the zygote has 46 chromosomes.
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Summarizing the Concepts
5.1 Maintaining the Chromosome Number
Each species has a characteristic number of chromosomes. The total
number is the diploid number, and half this number is the haploid
number. Among eukaryotes, cell division involves nuclear division
and division of the cytoplasm (cytokinesis).
Replication of DNA precedes cell division. The duplicated chro-
mosome is composed of two sister chromatids held together at a cen-
tromere. During mitosis the centromeres divide, and daughter
chromosomes go into each new nucleus.
The cell cycle has four stages. During the G
1
stage the organelles
increase in number; during the S stage, DNAreplication occurs; during
the G
2
stage, various proteins are synthesized; and during the M stage,
mitosis occurs. It is now known that regulation of the cell cycle
involves various proteins known as kinases and cyclins.
5.2 Mitosis in Detail
Mitosis has the following phases: prophasein early prophase chro-
mosomes have no particular arrangement, and in late prophase the
chromosomes are attached to spindle bers; metaphase, when the
chromosomes are aligned at the metaphase plate; anaphase, when the
chromatids separate, becoming daughter chromosomes that move
toward the poles; and telophase, when new nuclear envelopes form
around the daughter chromosomes and cytokinesis begins.
5.3 Reducing the Chromosome Number
Meiosis is found in any life cycle that involves sexual reproduction.
During meiosis I, homologues separate, and this leads to daughter cells
with half or the haploid number of homologous chromosomes.
Crossing-over and independent assortment of chromosomes during
meiosis I ensure genetic recombination in daughter cells. During meio-
sis II, chromatids separate, becoming daughter chromosomes that are
distributed to daughter nuclei. In some life cycles, the daughter cells
become gametes, and upon fertilization, the offspring have the diploid
number of chromosomes, the same as their parents.
5.4 Meiosis in Detail
Meiosis utilizes two nuclear divisions. During meiosis I, homologous
chromosomes undergo synapsis, and crossing-over between nonsister
chromatids occurs. When the homologous chromosomes separate dur-
ing meiosis I, each daughter nucleus receives one member from each
pair of chromosomes. Therefore, the daughter cells are haploid. Distri-
bution of daughter chromosomes derived from sister chromatids dur-
ing meiosis II then leads to a total of four new cells, each with the
haploid number of chromosomes.
5.5 Comparison of Meiosis with Mitosis
Figure 5.14 contrasts the phases of mitosis with the phases of meiosis.
5.6 The Human Life Cycle
The human life cycle involves both mitosis and meiosis. Mitosis ensures
that each somatic cell will have the diploid number of chromosomes.
Meiosis is a part of spermatogenesis and oogenesis. Spermatogen-
esis in males produces four viable sperm, while oogenesis in females
produces one egg and two polar bodies. Oogenesis does not go on to
completion unless a sperm fertilizes the developing egg.
Among sexually reproducing organisms, such as humans, meiosis
results in genetic recombination due to independent assortment of
homologous chromosomes and crossing-over. Fertilization also con-
tributes to genetic recombination.
Studying the Concepts
1. Describe the chromosome number using the terms diploid
and haploid. 84
2. Explain how mitosis maintains the chromosome number in
all the somatic cells of an individual. 84
3. Describe the cell cycle, including a description of interphase.
86
4. Describe the phases of animal mitosis, including in your
description the terms centrosome, nucleolus, spindle, and
cleavage furrow. 8891
100 Part 1 Cell Biology 5-18
C
loning is making exact multiple copies
of DNA or a cell or an organism. The
rst two procedures have been around for
some time. Through biotechnology, bacte-
ria produce cloned copies of human DNA.
When a single bacterium reproduces asex-
ually on a petri dish, a colony results. Each
member of the colony is a clone of the orig-
inal cell. Now for the rst time in our his-
tory, it is possible to produce a clone of a
vertebrate. No sperm and egg are
required. The DNA of an adult cell is
placed in an egg that undergoes develop-
ment to become an exact copy of the
organism that donated the DNA. Some
people fear that billionaires and celebrities
will hasten to make multiple copies of
themselves. Others feel that this is unlikely.
Rather, they fear a different type of
cloning.
Suppose it were possible to use the
DNA of a burn victim to produce embry-
onic cells that are cajoled to become skin
cells. These cells could be used to provide
grafts of brand new skin. Would this be a
proper use of cloning in humans?
Or suppose parents want to produce a
child free of a genetic disease. Scientists
produce a zygote through in vitro fertiliza-
tion, and then they clone the zygote to pro-
duce any number of cells. Genetic
engineering to correct the defect doesnt
work on all the cellsonly a few. They
implant just those few in the uterus where
development continues to term. Would
this be a proper use of cloning in humans?
Well, what if science progressed to
producing children with increased intelli-
gence or athletic prowess in the same
way? Would this be an acceptable use of
cloning in humans?
Questions
1. Presently, research in the cloning of
humans is banned. Should it be? Why or
why not?
2. Under what circumstances might cloning
in humans be acceptable? Explain.
3. Is cloning to produce improved breeds of
farm animals acceptable? Why or why
not?
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Go To Student OLC
9. At the metaphase plate during metaphase I of meiosis, there
are
a. single chromosomes.
b. unpaired duplicated chromosomes.
c. homologous pairs.
d. always twenty-three chromosomes.
10. Crossing-over occurs between
a. sister chromatids of the same chromosomes.
b. two different bivalents.
c. nonsister chromatids of a homologous pair.
d. two daughter nuclei.
11. Which of these is not a difference between spermatogenesis
and oogenesis in humans?
Spermatogenesis Oogenesis
occurs in males. occurs in females.
produces four sperm produces one egg
per meiosis. per meiosis.
produces haploid eggs produces diploid cells
always goes to does not always go to
completion completion
12. Label this diagram of a cell in early prophase of mitosis.
Chapter 5 Cell Division 101 5-19
5. Name two differences between plant cell mitosis and animal
cell mitosis. 91
6. Give an overview of meiosis and the manner in which it
reduces the chromosome number. 92
7. How does meiosis ensure genetic recombination in the
daughter cells? Explain in detail. 93
8. Describe the phases of meiosis I and meiosis II in detail. 94
9. Compare meiosis I and meiosis II to mitosis. 96
10. Explain why spermatogenesis results in four sperm and
oogenesis produces one mature egg. 9899
11. What are three events that ensure children have a different
combination of genes than their parents? 99
Testing Yourself
Choose the best answer for each question.
1. The cell cycle ensures that
a. the cell grows prior to cell division.
b. DNAreplicates prior to cell division.
c. the chromatids separate, becoming the daughter chromo-
somes.
d. the cytoplasm divides.
e. All of these are correct.
2. In human beings, mitosis is necessary to
a. growth and repair of tissues.
b. formation of the gametes.
c. maintaining the chromosome number in all body cells.
d. the death of unnecessary cells.
e. Both b and c are correct.
For questions 35 match the descriptions that follow to the terms
in the key.
Key:
a. centriole
b. chromatid
c. chromosome
d. centromere
3. point of attachment for sister chromatids
4. found at a pole in the center of an aster
5. coiled and condensed chromatin
6. If a parent cell has fourteen chromosomes prior to mitosis,
how many chromosomes will the daughter cells have?
a. twenty-eight
b. fourteen
c. seven
d. any number between seven and twenty-eight
7. In which phase of mitosis are chromosomes moving toward
the poles?
a. prophase
b. metaphase
c. anaphase
d. telophase
e. Both b and c are correct.
8. If a parent cell has twelve chromosomes, then the daughter
cells following meiosis will have
a. twelve chromosomes.
b. twenty-four chromosomes.
c. six chromosomes.
d. Any one of these could be correct.
a.
b.
c.
d.
13. Which of these drawings represents metaphase I of meiosis?
How do you know?
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102 Part 1 Cell Biology 5-20
Match the terms to these denitions:
a. Production of sperm in males by the process of
meiosis and maturation.
b. In oogenesis, a nonfunctional product; two to
three meiotic products are of this type.
c. In oogenesis, the functional product of meiosis I;
becomes the egg.
d. Pairing of homologous chromosomes during
meiosis I.
e. Production of eggs in females by the process of
meiosis and maturation.
Understanding the Terms
anaphase 89
aster 88
cell cycle 86
cell plate 90
centromere 84
chromatin 84
chromosome 84
cleavage furrow 91
crossing-over 93
cyclin 86
cytokinesis 84
daughter chromosomes 84
diploid (2n) 84
fertilization 92
gamete 92
haploid (n) 84
homologous chromosome 92
homologue 92
independent assortment 93
interkinesis 94
interphase 86
kinase 86
meiosis 92
metaphase 89
mitosis 84
oogenesis 98
polar body 98
prophase 88
secondary oocyte 98
sister chromatid 84
somatic cell 84
spermatogenesis 98
spindle 88
synapsis 92
telophase 89
zygote 98
Thinking Scientically
1. Concerning the genetic material:
a. Which formchromatin or chromosomeswould you
expect to be metabolically active? Why? (page 84)
b. The genes are part of the chromosomes. Ordinarily, a per-
son inherits two copies of a geneone from the mother
and one from the father. Does it seem reasonable that the
two copies of the gene might be different forms of the
gene?
c. The genes direct protein (enzyme) synthesis. Why do you
suppose a person with three chromosomes of the same
kind might suffer from various disorders?
d. Not all genes are active in all cells. For example, why
might a defective gene not adversely affect the maturation
of a sperm or an egg?
2. Concerning cell division (page 96):
a. The drug colchicine prevents cell division from nishing.
What part of a dividing cell do you suppose it disrupts?
b. Asexual reproduction ordinarily does not produce genetic
variation. Why not?
c. Sexual reproduction does produce genetic variation. Why?
d. Would you expect sexual reproduction to aid the evolu-
tionary process? Why?
Using Technology
Your study of cell division is supported by these available
technologies:
Essential Study Partner CD-ROM
Cells Cell Division
Visit the Mader web site for related ESP activities.
Exploring the Internet
The Mader Home Page provides resources and tools as
you study this chapter.
http://www.mhhe.com/biosci/genbio/mader
Life Science Animations 3DVideo
10 Mitosis
11 Meiosis
12 Crossing Over
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