The Cell Cycle and How Cells Divide

Phases of the Cell Cycle

The cell cycle consists of
Interphase normal cell activity
The mitotic phase cell divsion
INTERPHASE

Growth
G1
Ce
ll
Di
vs
io
n

(DNA synthesis)
Growth
G2

Functions of Cell Division

100 m

(a) Reproduction. An amoeba,

a single-celled eukaryote, is
dividing into two cells. Each
new cell will be an individual
organism (LM).

200 m

20 m

(b) Growth and development.

(c) Tissue renewal. These dividing
This micrograph shows a
bone marrow cells (arrow) will
sand dollar embryo shortly after
give rise to new blood cells (LM).
the fertilized egg divided, forming
two cells (LM).

Cell Division

An integral part of the cell cycle

Results in genetically identical daughter cells
Cells duplicate their genetic material
Before they divide, ensuring that each daughter
cell receives an exact copy of the genetic
material, DNA

DNA

Genetic information - genome

Packaged into chromosomes

Figure 12.3
50 m

DNA And Chromosomes

An average eukaryotic cell has about 1,000

times more DNA then an average
prokaryotic cell.
The DNA in a eukaryotic cell is organized
into several linear chromosomes, whose
organization is much more complex than the
single, circular DNA molecule in a
prokaryotic cell

Chromosomes

All eukaryotic cells store genetic information

in chromosomes.
Most eukaryotes have between 10 and 50
chromosomes in their body cells.
Human cells have 46 chromosomes.
23 nearly-identical pairs

Structure of Chromosomes

Chromosomes are composed of a

complex of DNA and protein called
chromatin that condenses during cell
division
DNA exists as a single, long, doublestranded fiber extending chromosomes
entire length.
Each unduplicated chromosome contains
one DNA molecule, which may be
several inches long
8

Structure of Chromosomes

Every 200 nucleotide pairs, the DNA wraps twice around a

group of 8 histone proteins to form a nucleosome.
Higher order coiling and supercoiling also help condense
and package the chromatin inside the nucleus:

Structure of Chromosomes

The degree of coiling can vary in different

regions of the chromatin:
Heterochromatin refers to highly coiled
regions where genes arent expressed.
Euchromatin refers to loosely coiled regions
where genes can be expressed.

10

Structure of Chromosomes

Prior to cell division each

chromosome duplicates
itself.
During this time, only the
heterochromatin is visible, as
dense granules inside the
nucleus.
There is also a dense area of
RNA production called the
nucleolus:
11

Karyotype

An ordered, visual representation of the chromosomes in a cell

Chromosomes are photographed when they are highly condensed, then photos
of the individual chromosomes are arranged in order of decreasing size:
In humans each somatic cell has 46 chromosomes, made up of two sets, one
set of chromosomes comes from each parent

Pair of homologous
chromosomes

5 m

Centromere
Sister
chromatids

12

Chromosomes

Non-homologous chromosomes
Look different
Control different traits
Sex chromosomes
Are distinct from each other in their
characteristics
Are represented as X and Y
Determine the sex of the individual, XX being
female, XY being male
In a diploid cell, the chromosomes occur in pairs.
The 2 members of each pair are called
homologous chromosomes or homologues.
13

Chromosomes

A diploid cell has two sets of each of its chromosomes

A human has 46 chromosomes (2n = 46)
In a cell in which DNA synthesis has occurred all the chromosomes are
duplicated and thus each consists of two identical sister chromatids
Maternal set of
chromosomes (n = 3)
2n = 6
Paternal set of
chromosomes (n = 3)

Two sister chromatids

of one replicated
chromosome
Centromere

Two nonsister
chromatids in
a homologous pair

Pair of homologous
chromosomes
(one from each set)

14

Homologues

Homologous chromosomes:

Look the same

Control the same traits

May code for different forms of each trait

Independent origin - each one was inherited

from a different parent

15

Chromosome Duplication

In preparation for cell division, DNA is replicated and the chromosomes condense

Each duplicated chromosome has two sister chromatids, which separate during cell
division
0.5 m
A eukaryotic cell has multiple
chromosomes, one of which is
represented here. Before
duplication, each chromosome
has a single DNA molecule.

Once duplicated, a chromosome

consists of two sister chromatids
connected at the centromere. Each
chromatid contains a copy of the
DNA molecule.

Mechanical processes separate

the sister chromatids into two
chromosomes and distribute
them to two daughter cells.

Chromosome
duplication
(including DNA
synthesis)
Centromere

Separation
of sister
chromatids

Centrometers

Sister
chromatids

Sister chromatids

16

Chromosome Duplication

Because of duplication, each condensed chromosome

consists of 2 identical chromatids joined by a centromere.
Each duplicated chromosome contains 2 identical DNA
molecules (unless a mutation occurred), one in each
chromatid:
Non-sister
chromatids

Centromere

Duplication

Sister
chromatids
Two unduplicated
chromosomes

Sister
chromatids

Two duplicated chromosomes

Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

17

Structure of Chromosomes

The centromere is a constricted region of the chromosome containing a

specific DNA sequence, to which is bound 2 discs of protein called
kinetochores.
Kinetochores serve as points of attachment for microtubules that move
the chromosomes during cell division:
Metaphase chromosome

Centromere
region of
chromosome

Kinetochore

Kinetochore
microtubules

Sister Chromatids
18
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Structure of Chromosomes

Diploid - A cell possessing two copies of each chromosome

(human body cells).
Homologous chromosomes are made up of sister
chromatids joined at the centromere.
Haploid - A cell possessing a single copy of each
chromosome (human sex cells).

19

Phases of the Cell Cycle

Interphase

G1 - primary growth

S - genome replicated
G2 - secondary growth

M - mitosis
C - cytokinesis

20

Interphase

G1 - Cells undergo majority of growth

S - Each chromosome replicates (Synthesizes) to
produce sister chromatids
Attached at centromere
Contains attachment site (kinetochore)

G2 - Chromosomes condense - Assemble

machinery for division such as centrioles

21

Mitosis

Some haploid & diploid cells divide by mitosis.

Each new cell receives one copy of every
chromosome that was present in the original cell.
Produces 2 new cells that are both genetically
identical to the original cell.
DNA
duplication
during
interphase

Mitosis

Diploid Cell

22

Mitotic Division of an Animal Cell

G2 OF INTERPHASE

Centrosomes
(with centriole pairs)

Nucleolus

Chromatin
(duplicated)

Nuclear
Plasma
envelope membrane

PROPHASE

Early mitotic
spindle

Aster
Centromere

Chromosome, consisting
of two sister chromatids

PROMETAPHASE

Fragments
of nuclear
envelope

Kinetochore
Nonkinetochore
microtubules

Kinetochore
microtubule
23

Mitotic Division of an Animal Cell

METAPHASE

ANAPHASE

Metaphase
plate

Spindle

Centrosome at Daughter
one spindle pole chromosomes

TELOPHASE AND CYTOKINESIS

Cleavage
furrow

Nucleolus
forming

Nuclear
envelope
forming
24

G2 of Interphase
A nuclear envelope bounds
the nucleus.
The nucleus contains one or
more nucleoli (singular,
nucleolus).
Two centrosomes have
formed by replication of a
single centrosome.
In animal cells, each
centrosome features two
centrioles.
Chromosomes, duplicated
during S phase, cannot be
seen individually because
they have not yet condensed.
The light micrographs show dividing lung cells
from a newt, which has 22 chromosomes in
its somatic cells (chromosomes appear blue,
microtubules green, intermediate filaments
red). For simplicity, the drawings show only
four chromosomes.

G2 OF INTERPHASE

Centrosomes
(with centriole pairs)

Nucleolus

Chromatin
(duplicated)

Nuclear
Plasma
envelope membrane
25

Prophase
The chromatin fibers become
more tightly coiled, condensing
into discrete chromosomes
observable with a light
microscope.
The nucleoli disappear.
Each duplicated chromosome
appears as two identical sister
chromatids joined together.
The mitotic spindle begins to form.
It is composed of the centrosomes
and the microtubules that extend
from them. The radial arrays of
shorter microtubules that extend
from the centrosomes are called
asters (stars).
The centrosomes move away from
each other, apparently propelled
by the lengthening microtubules
between them.

PROPHASE

Early mitotic
spindle

Aster
Centromere

Chromosome, consisting
of two sister chromatids
26

Metaphase
Metaphase is the longest stage of
mitosis, lasting about 20 minutes.
The centrosomes are now at
opposite ends of the cell.
The chromosomes convene on the
metaphase plate, an imaginary
plane that is equidistant between
the spindles two poles. The
chromosomes centromeres lie on
the metaphase plate.
For each chromosome, the
kinetochores of the sister
chromatids are attached to
kinetochore microtubules coming
from opposite poles.
The entire apparatus of
microtubules is called the spindle
because of its shape.

METAPHASE
Metaphase
plate

Spindle

Centrosome at
one spindle pole
27

The Mitotic Spindle

The spindle includes the centrosomes, the spindle

microtubules, and the asters
The apparatus of microtubules controls
chromosome movement during mitosis
The centrosome replicates, forming two
centrosomes that migrate to opposite ends of the
cell
Assembly of spindle microtubules begins in the
centrosome, the microtubule organizing center
An aster (a radial array of short microtubules)
extends from each centrosome
28

The Mitotic Spindle

Some spindle microtubules attach to the kinetochores of

chromosomes and move the chromosomes to the metaphase
plate
In anaphase, sister chromatids separate and move along the
kinetochore microtubules toward opposite ends of the cell
Aster
Microtubules

Sister
chromatids

Chromosomes

Centrosome
Metaphase
plate
Kinetochores

Centrosome

1 m

Overlapping
nonkinetochore
microtubules

Kinetochore
microtubules

0.5 m

29

Anaphase
Anaphase is the shortest stage of
mitosis, lasting only a few minutes.
Anaphase begins when the two sister
chromatids of each pair suddenly part.
Each chromatid thus becomes a fullfledged chromosome.
The two liberated chromosomes begin
moving toward opposite ends of the cell,
as their kinetochore microtubules
shorten. Because these microtubules are
attached at the centromere region, the
chromosomes move centromere first (at
about 1 m/min).
The cell elongates as the
nonkinetochore microtubules lengthen.
By the end of anaphase, the two ends of
the cell have equivalentand
completecollections of chromosomes.

ANAPHASE

Daughter
chromosomes
30

Telophase
Two daughter nuclei begin to
form in the cell.
Nuclear envelopes arise from
the fragments of the parent
cells nuclear envelope and
other portions of the
endomembrane system.
The chromosomes become
less condensed.
Mitosis, the division of one
nucleus into two genetically
identical nuclei, is now
complete.

TELOPHASE AND CYTOKINESIS

Cleavage
furrow

Nucleolus
forming

Nuclear
envelope
forming

31

Mitosis in a plant cell

Chromatine
Nucleus
Nucleolus condensing

1 Prophase.
The chromatin
is condensing.
The nucleolus is
beginning to
disappear.
Although not
yet visible
in the micrograph,
the mitotic spindle is
staring to from.

Chromosome

Metaphase. The
2 Prometaphase.
3
4
spindle is complete,
We now see discrete
and the chromosomes,
chromosomes; each
attached to microtubules
consists of two
at their kinetochores,
identical sister
are all at the metaphase
chromatids. Later
plate.
in prometaphase, the
nuclear envelop will
fragment.

5
Anaphase. The
chromatids of each
chromosome have
separated, and the
daughter chromosomes
are moving to the ends
of cell as their
kinetochore
microtubles shorten.

Telophase. Daughter
nuclei are forming.
Meanwhile, cytokinesis
has started: The cell
plate, which will
divided the cytoplasm
in two, is growing
toward the perimeter
of the parent cell.

32

Cytokinesis

Cleavage of cell into two

halves
Animal cells
Constriction belt of
actin filaments
Plant cells
Cell plate
Fungi and protists
Mitosis occurs
within the nucleus
33

Cytokinesis In Animal And Plant Cells

Cleavage furrow

Contractile ring of
microfilaments

100 m

Vesicles
forming
cell plate

Wall of
patent cell

1 m

Cell plate

New cell wall

Daughter cells
Daughter cells

(a) Cleavage of an animal cell (SEM)

(b) Cell plate formation in a plant cell (SEM)

34

35

Meiosis and Sexual Life Cycles

Living organisms are distinguished by their ability to

reproduce their own kind
Heredity
Is the transmission of traits from one generation to the
next
Variation
Shows that offspring differ somewhat in appearance
from parents and siblings

36

Inheritance of Genes

Genes are segments of DNA, units of heredity

Offspring acquire genes from parents by
inheriting chromosomes
Genetics is the scientific study of heredity and
hereditary variation

37

Inheritance of Genes

Each gene in an organisms DNA has a

specific locus on a certain chromosome
We inherit one set of chromosomes from our
mother and one set from our father
Two parents give rise to offspring that have
unique combinations of genes inherited from
the two parents - sexual reproduction

38

Asexual Reproduction

In asexual reproduction, one parent

produces genetically identical offspring by
mitosis

Parent
Bud

Figure 13.2

0.5 mm
39

Sexual Reproduction

Fertilization and meiosis alternate in sexual life cycles

A life cycle is the generation-to-generation sequence of
stages in the reproductive history of an organism
Key
Haploid
Diploid
n

Gametes
n

MEIOSIS

2n
Diploid
multicellular
organism

FERTILIZATION

Zygote

2n

Mitosis
(a) Animals

40

Sex Cells - Gametes

Unlike somatic cells, sperm and egg cells

are haploid cells, containing only one set of
chromosomes
At sexual maturity the ovaries and testes
produce haploid gametes by meiosis

41

Sexual Reproduction - The Human Life Cycle

During fertilization,
sperm and ovum fuse
forming a diploid
zygote
The zygote develops
into an adult organism

Haploid gametes (n = 23)

Haploid (n)
Diploid (2n)

Ovum (n)

Sperm
Cell (n)
FERTILIZATION

MEIOSIS

Ovary

Testis

Diploid
zygote
(2n = 46)

Mitosis and
development
Multicellular diploid
adults (2n = 46)

42

Meiosis

Reduces the chromosome number such that

each daughter
Cell has a haploid set of chromosomes
Ensures that the next generation will have:
Diploid number of chromosome
Exchange of genetic information
(combination of traits
that differs from that of either parent)
43

Meiosis

Only diploid cells can divide by meiosis.

Prior to meiosis I, DNA replication occurs.
During meiosis, there will be two nuclear divisions, and the result will be
four haploid nuclei.
No replication of DNA occurs between meiosis I and meiosis II.

44

Meiosis
Interphase

Meiosis reduces the

number of chromosome
sets from diploid to
haploid
Meiosis takes place in
two sets of divisions

Meiosis I reduces the

number of chromosomes
from diploid to haploid
Meiosis II produces four
haploid daughter cells
Figure 13.7

Homologous pair
of chromosomes
in diploid parent cell

Chromosomes
replicate
Homologous pair of replicated chromosomes

Sister
chromatids

Diploid cell with

replicated
chromosomes

Meiosis I

1 Homologous
chromosomes
separate

Haploid cells with

replicated chromosomes
Meiosis II
2 Sister chromatids
separate

Haploid cells with unreplicated chromosomes

45

Meiosis Phases

Meiosis involves the same four phases seen in

mitosis
prophase
metaphase
anaphase
telophase
They are repeated during both meiosis I and
meiosis II.
The period of time between meiosis I and meiosis
II is called interkinesis.
No replication of DNA occurs during interkinesis
because the DNA is already duplicated.
46

Prophase I

Prophase I occupies more than 90% of the time required for meiosis
Chromosomes begin to condense
In synapsis, the 2 members of each homologous pair of chromosomes line
up side-by-side, aligned gene by gene, to form a tetrad consisting of 4
chromatids
During synapsis, sometimes there is an exchange of homologous parts
between non-sister chromatids. This exchange is called crossing over
Each tetrad usually has one or more chiasmata, X-shaped regions where
crossing over occurred

Nonsister
chromatids

Prophase I
of meiosis

Tetrad

Chiasma,
site of
crossing
over

47

Metaphase I

At metaphase I, tetrads line up at the metaphase plate, with one

chromosome facing each pole
Microtubules from one pole are attached to the kinetochore of one
chromosome of each tetrad
Microtubules from the other pole are attached to the kinetochore of the
other chromosome
PROPHASE I
Sister
chromatids

Tetrad

METAPHASE I

ANAPHASE I
Sister chromatids
remain attached

Centromere
(with kinetochore)

Chiasmata

Metaphase
plate

Spindle

Microtubule
attached to
kinetochore

Homologous chromosomes
(red and blue) pair and
exchange segments; 2n = 6

Homologous
chromosomes
separate

Tetrads line up

Pairs of homologous
chromosomes split up

48

Anaphase I

In anaphase I, pairs of homologous chromosomes separate

One chromosome moves toward each pole, guided by the
spindle apparatus
Sister chromatids remain attached at the centromere and
move as one unit toward the pole
PROPHASE I
Sister
chromatids

Tetrad

METAPHASE I

ANAPHASE I
Sister chromatids
remain attached

Centromere
(with kinetochore)

Chiasmata

Metaphase
plate

Spindle

Microtubule
attached to
kinetochore

Homologous chromosomes
(red and blue) pair and
exchange segments; 2n = 6

Homologous
chromosomes
separate

Tetrads line up

Pairs of homologous
chromosomes split up

49

Telophase I and Cytokinesis

In the beginning of telophase I, each half of the

cell has a haploid set of chromosomes; each
chromosome still consists of two sister chromatids
Cytokinesis usually occurs simultaneously, forming
two haploid daughter cells
In animal cells, a cleavage furrow forms; in plant
cells, a cell plate forms
No chromosome replication occurs between the
end of meiosis I and the beginning of meiosis II
because the chromosomes are already replicated

50

Prophase II

Meiosis II is very similar to mitosis

In prophase II, a spindle apparatus forms
In late prophase II, chromosomes (each still composed of
two chromatids) move toward the metaphase plate
TELOPHASE I AND
CYTOKINESIS

PROPHASE II

Cleavage
furrow

METAPHASE II

ANAPHASE II

Sister chromatids
separate

TELOPHASE II AND
CYTOKINESIS

Haploid daughter cells

forming

51

Metaphase II

At metaphase II, the sister chromatids are at the metaphase plate

Because of crossing over in meiosis I, the two sister chromatids of each
chromosome are no longer genetically identical
The kinetochores of sister chromatids attach to microtubules extending
from opposite poles
TELOPHASE I AND
CYTOKINESIS

PROPHASE II

Cleavage
furrow

METAPHASE II

ANAPHASE II

Sister chromatids
separate

TELOPHASE II AND
CYTOKINESIS

Haploid daughter cells

forming

52

Anaphase II

At anaphase II, the sister chromatids separate

The sister chromatids of each chromosome now move as
two newly individual chromosomes toward opposite poles
TELOPHASE I AND
CYTOKINESIS

PROPHASE II

Cleavage
furrow

METAPHASE II

ANAPHASE II

Sister chromatids
separate

TELOPHASE II AND
CYTOKINESIS

Haploid daughter cells

forming

53

Telophase II and Cytokinesis

In telophase II, the chromosomes arrive at opposite poles

Nuclei form, and the chromosomes begin decondensing
Cytokinesis separates the cytoplasm
At the end of meiosis, there are four daughter cells, each with a
haploid set of unreplicated chromosomes
Each daughter cell is genetically distinct from the others and from the
parent cell
TELOPHASE I AND
CYTOKINESIS

PROPHASE II

Cleavage
furrow

METAPHASE II

ANAPHASE II

Sister chromatids
separate

TELOPHASE II AND
CYTOKINESIS

Haploid daughter cells

forming

54

A Comparison of Mitosis and Meiosis

Mitosis conserves the number of chromosome

sets, producing cells that are genetically identical
to the parent cell
Meiosis reduces the number of chromosomes sets
from two (diploid) to one (haploid), producing cells
that differ genetically from each other and from the
parent cell
The mechanism for separating sister chromatids is
virtually identical in meiosis II and mitosis

55

A Comparison of Mitosis and Meiosis

Three events are unique to meiosis, and all three

occur in meiosis l:

Synapsis and crossing over in prophase I:

Homologous chromosomes physically connect and
exchange genetic information
At the metaphase plate, there are paired homologous
chromosomes (tetrads), instead of individual replicated
chromosomes
At anaphase I of meiosis, homologous pairs move
toward opposite poles of the cell. In anaphase II of
meiosis, the sister chromatids separate

56

A Comparison Of Mitosis And Meiosis

MITOSIS

MEIOSIS
Chiasma (site of
crossing over)

Parent cell
(before chromosome replication)

MEIOSIS I

Prophase I

Prophase

Chromosome
replication

Duplicated chromosome
(two sister chromatids)

Chromosome
replication

Tetrad formed by
synapsis of homologous
chromosomes

2n = 6

Chromosomes
positioned at the
metaphase plate

Metaphase

Sister chromatids
separate during
anaphase

Anaphase
Telophase

2n

Tetrads
positioned at the
metaphase plate

Homologues
separate
during
anaphase I;
sister
chromatids
remain together

Metaphase I

Anaphase I
Telophase I
Haploid
n=3

Daughter
cells of
meiosis I

2n

MEIOSIS II

Daughter cells
of mitosis
n

Daughter cells of meiosis II

Sister chromatids separate during anaphase II

57

Comparison

Meiosis
DNA duplication
followed by 2 cell
divisions
Sysnapsis
Crossing-over
One diploid cell
produces 4
haploid cells
Each new cell
has a unique
combination of
genes

Mitosis
Homologous
chromosomes do not
pair up
No genetic exchange
between homologous
chromosomes
One diploid cell
produces 2 diploid
cells or one haploid
cell produces 2
haploid cells
New cells are
genetically identical to
original cell (except for
mutation)

58

Sexual Reproduction - The Human Life Cycle

During fertilization,
sperm and ovum fuse
forming a diploid
zygote
The zygote develops
into an adult organism

Haploid gametes (n = 23)

Haploid (n)
Diploid (2n)

Ovum (n)

Sperm
Cell (n)
FERTILIZATION

MEIOSIS

Ovary

Testis

Diploid
zygote
(2n = 46)

Mitosis and
development
Multicellular diploid
adults (2n = 46)

59

Spermatocytes to Spermatids

Primary spermatocytes undergo meiosis I, forming

two haploid cells called secondary spermatocytes
Secondary spermatocytes undergo meiosis II and
their daughter cells are called spermatids
Spermatids are small round cells seen close to the
lumen of the tubule
Late in spermatogenesis, spermatids are nonmotile
Spermiogenesis spermatids lose excess
cytoplasm and form a tail, becoming motile sperm

60

Spermatogenesis

Figure 27.8b, c

61

Oogenesis

Production of female sex cells by meiosis

In the fetal period, oogonia (2n ovarian stem cells) multiply by
mitosis and store nutrients
Primordial follicles appear as oogonia are transformed into
primary oocytes
Primary oocytes begin meiosis but stall in prophase I
From puberty, each month one activated primary oocyte
completes meiosis one to produce two haploid cells
The first polar body
The secondary oocyte
The secondary oocyte arrests in metaphase II and is ovulated
If penetrated by sperm the second oocyte completes meiosis
II, yielding:
One large ovum (the functional gamete)
A tiny second polar body
62

Oogenesis

63