The Cell Cycle and How Cells Divide

1
 Phases of the Cell Cycle
• The cell cycle consists of
– Interphase – normal cell activity

– The mitotic phase – cell divsion

INTERPHASE

Growth
G1 (DNA synthesis)

Growth
G2

2
 Functions of Cell Division

100 µm 200 µm 20 µm

(a) Reproduction. An amoeba,
a single-celled eukaryote, is
dividing into two cells. Each
new cell will be an individual
organism (LM).
(b) Growth and development.
This micrograph shows a
sand dollar embryo shortly after
the fertilized egg divided, forming
two cells (LM).
(c) Tissue renewal. These dividing
bone marrow cells (arrow) will
give rise to new blood cells (LM).

3
 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

4
 DNA
• Genetic information - genome
• Packaged into chromosomes

Figure 12.3
50 µm

5
 DNA And Chromosomes

• An average eukaryotic cell has about 1,000

times more DNA than 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

6
 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

7
 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, double-
stranded fiber extending chromosome’s
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:

9
 Structure of Chromosomes
The degree of coiling can vary in different
regions of the chromatin:
Heterochromatin refers to highly coiled
regions where genes aren’t 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 µm

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 Pair of homologous

chromatids in chromosomes
a homologous pair (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 µm
A eukaryotic cell has multiple
chromosomes, one of which is
represented here. Before
duplication, each chromosome
has a single DNA molecule. Chromosome
duplication
(including DNA
synthesis)
Once duplicated, a chromosome
Centromere
consists of two sister chromatids
connected at the centromere. Each
chromatid contains a copy of the
DNA molecule.

Sister
Separation chromatids
of sister
chromatids
Mechanical processes separate
the sister chromatids into two
chromosomes and distribute
them to two daughter cells.
16
Centrometers Sister chromatids
 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 Sister
chromatids chromatids
Two unduplicated
chromosomes Two duplicated chromosomes
17
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
 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 PROPHASE PROMETAPHASE

Centrosomes Aster Fragments
(with centriole pairs) Chromatin Early mitotic Kinetochore
spindle Centromere of nuclear
(duplicated) Nonkinetochore
envelope
microtubules

Nucleolus Nuclear Plasma Chromosome, consisting Kinetochore

envelope membrane of two sister chromatids microtubule
23
 Mitotic Division of an Animal Cell

METAPHASE ANAPHASE TELOPHASE AND CYTOKINESIS

Metaphase
plate Cleavage Nucleolus
furrow forming

Nuclear
envelope
Spindle Centrosome at Daughter forming
one spindle pole chromosomes
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 G2 OF INTERPHASE
centrosome features two
Centrosomes
centrioles. (with centriole pairs) Chromatin
• Chromosomes, duplicated (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.
Nucleolus 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
PROPHASE
chromatids joined together.
• The mitotic spindle begins to form. Early mitotic
Aster
It is composed of the centrosomes spindle Centromere
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.
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
METAPHASE
plane that is equidistant between
the spindle’s two poles. The Metaphase
plate
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
Spindle Centrosome at
because of its shape. 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
Sister Centrosome
Microtubules Chromosomes chromatids Metaphase
plate
Kineto-
chores

Overlapping
nonkinetochore Kinetochore
microtubules microtubules
Centrosome 1 µm 0.5 µm
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 full-
fledged chromosome.
• The two liberated chromosomes begin
moving toward opposite ends of the cell, ANAPHASE
as their kinetochore microtubules
shorten. Because these microtubules are
attached at the centromere region, the
chromosomes move centromere first (at
about 1 µm/min).
• The cell elongates as the
nonkinetochore microtubules lengthen.
• By the end of anaphase, the two ends of
the cell have equivalent—and
complete—collections of chromosomes.
Daughter
chromosomes

30
Telophase
• Two daughter nuclei begin to
form in the cell.
• Nuclear envelopes arise from
the fragments of the parent
cell’s nuclear envelope and TELOPHASE AND CYTOKINESIS
other portions of the
endomembrane system. Cleavage Nucleolus
furrow
• The chromosomes become forming

less condensed.
• Mitosis, the division of one
nucleus into two genetically
identical nuclei, is now
Nuclear
complete. envelope
forming

31
 Mitosis in a plant cell
Nucleus Chromatine Chromosome
Nucleolus condensing

Metaphase. The
1 Prophase. 2 Prometaphase. 3 4 Anaphase. The 5 Telophase. Daughter
spindle is complete,
The chromatin We now see discrete chromatids of each nuclei are forming.
and the chromosomes,
is condensing. chromosomes; each chromosome have Meanwhile, cytokinesis
attached to microtubules
The nucleolus is consists of two separated, and the has started: The cell
at their kinetochores,
beginning to identical sister daughter chromosomes plate, which will
are all at the metaphase
disappear. chromatids. Later are moving to the ends divided the cytoplasm
plate.
Although not in prometaphase, the of cell as their in two, is growing
yet visible nuclear envelop will kinetochore toward the perimeter
in the micrograph, fragment. microtubles shorten. of the parent cell.
the mitotic spindle is
staring to from.

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

100 µm
Cleavage furrow Vesicles Wall of 1 µm
forming patent cell Cell plate
cell plate New cell wall

Contractile ring of Daughter cells

microfilaments
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 organism’s 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 n
Gametes
n

MEIOSIS FERTILIZATION

Zygote
2n 2n

Diploid Mitosis
multicellular
organism
(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
Haploid gametes (n = 23)
Haploid (n)
• During fertilization, Diploid (2n)
Ovum (n)

sperm and ovum fuse

forming a diploid Sperm
Cell (n)
zygote
MEIOSIS FERTILIZATION
• The zygote develops
into an adult organism
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
Homologous pair

• Meiosis reduces the of chromosomes

in diploid parent cell

number of chromosome
sets from diploid to Chromosomes

haploid Homologous pair of replicated chromosomes

replicate

• Meiosis takes place in Sister

two sets of divisions chromatids Diploid cell with

replicated
chromosomes

– Meiosis I reduces the Meiosis I

number of chromosomes 1 Homologous

from diploid to haploid chromosomes

separate

Haploid cells with

– Meiosis II produces four Meiosis II

replicated chromosomes

haploid daughter cells 2 Sister chromatids

separate

Haploid cells with unreplicated chromosomes

Figure 13.7 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

Prophase I Nonsister
of meiosis chromatids

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 METAPHASE I ANAPHASE I

Centromere Sister chromatids
(with kinetochore) remain attached
Sister Chiasmata
chromatids Metaphase
Spindle plate

Tetrad Microtubule Homologous

attached to chromosomes
kinetochore separate
Homologous chromosomes Tetrads line up Pairs of homologous
(red and blue) pair and chromosomes split up 48
exchange segments; 2n = 6
 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 METAPHASE I ANAPHASE I
Centromere Sister chromatids
(with kinetochore) remain attached
Sister Chiasmata
chromatids Metaphase
Spindle plate

Tetrad Microtubule Homologous

attached to chromosomes
kinetochore separate
Homologous chromosomes Tetrads line up Pairs of homologous
(red and blue) pair and chromosomes split up 49
exchange segments; 2n = 6
 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 PROPHASE II TELOPHASE II AND
METAPHASE II ANAPHASE II
CYTOKINESIS CYTOKINESIS

Cleavage Haploid daughter cells

furrow Sister chromatids forming
separate

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 PROPHASE II TELOPHASE II AND

METAPHASE II ANAPHASE II
CYTOKINESIS CYTOKINESIS

Cleavage Haploid daughter cells

furrow Sister chromatids forming
separate

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 PROPHASE II TELOPHASE II AND

METAPHASE II ANAPHASE II
CYTOKINESIS CYTOKINESIS

Cleavage Haploid daughter cells

furrow Sister chromatids forming
separate

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 PROPHASE II TELOPHASE II AND
METAPHASE II ANAPHASE II
CYTOKINESIS CYTOKINESIS

Cleavage Haploid daughter cells

furrow Sister chromatids forming
separate

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
Parent cell Chiasma (site of
MEIOSIS I
(before chromosome replication) crossing over)

Prophase Prophase I
Chromosome Chromosome
replication replication Tetrad formed by
Duplicated chromosome 2n = 6 synapsis of homologous
(two sister chromatids) chromosomes

Chromosomes Tetrads
positioned at the positioned at the Metaphase I
Metaphase
metaphase plate metaphase plate

Anaphase Sister chromatids Homologues

Telophase separate during Anaphase I
separate
anaphase Telophase I
during
anaphase I;
sister Haploid
chromatids n=3
Daughter
remain together
cells of
meiosis I
2n 2n
Daughter cells MEIOSIS II
of mitosis

n n n n
Daughter cells of meiosis II

Sister chromatids separate during anaphase II

57
 Comparison
• Mitosis
• Meiosis
• Homologous
• DNA duplication
chromosomes do not
followed by 2 cell
pair up
divisions
• No genetic exchange
• Sysnapsis
between homologous
• Crossing-over chromosomes
• One diploid cell • One diploid cell
produces 4 produces 2 diploid
haploid cells cells or one haploid
• Each new cell cell produces 2
has a unique haploid cells
combination of • New cells are
genes genetically identical to
original cell (except for
mutation)

58
 Sexual Reproduction - The Human Life Cycle
Haploid gametes (n = 23)
Haploid (n)
• During fertilization, Diploid (2n)
Ovum (n)

sperm and ovum fuse

forming a diploid Sperm
Cell (n)
zygote
MEIOSIS FERTILIZATION
• The zygote develops
into an adult organism
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