BIOLOGY
tenth edition
Chapter 10
Chromosomes, Mitosis,
and Meiosis
© Cengage Learning 2015 SOLOMON • MARTIN • MARTIN •
BERG
Deoxyribonucleic Acid (DNA)
• Cells contain DNA: coded genetic
information
– Genes control a cell’s activities and are
passed on to descendants
• When a cell divides, DNA
is replicated and copies
are transmitted to each
daughter cell
© Cengage Learning 2015
Chromosomes, Mitosis, and Meiosis
• Each DNA molecule is packaged with
proteins to form a chromosome
– Mitosis: a parent cell transmits one copy of
every chromosome to each of its two
daughter cells
– Meiosis: in organisms that reproduce sexually,
chromosome number is reduced in half to
produce eggs and sperm, then the original
number is restored in the zygote
© Cengage Learning 2015
10.1 Eukaryotic Chromosomes
• When cell is not dividing, chromosomes
are present in extended, partially
unraveled form
• During cell division, chromatin fibers
condense and chromosomes
become visible as
distinct structures
© Cengage Learning 2015
Histones and Nucleosomes
• In eukaryotic cells, DNA is packed in
multiple chromosomes
DNA wound around Histone tails
a cluster of histone
molecules
Linker DNA
Nucleosome
(10 nm diameter)
© Cengage Learning 2015
DNA Released From Histone Proteins
• Histones are an DNA
important part of the loops
regulation of gene
expression – Scaffolding
whether genes are proteins
turned off or on
© Cengage Learning 2015
Levels of Chromosome Structure
• Nucleosomes are 10 nm in diameter
• Histone H1 associates with linker DNA,
packing adjacent nucleosomes together in
a 30 nm chromatin fiber
• Chromatin fibers form large, coiled loops
held together by scaffolding proteins
• Condensin binds to DNA and wraps it into
coiled loops that are compacted into a
chromosome
© Cengage Learning 2015
ANIMATION: Chromosome Structural
Organization
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Chromosome Number and Content
• Every species has a characteristic number
of chromosomes in the nucleus of each
somatic cell
– Humans: 46 chromosomes
– Roundworm: 2 chromosomes
– Some ferns have more than 1000
chromosomes
• It is the information specified in the genes
that makes each species unique
© Cengage Learning 2015
10.2 The Cell Cycle and Mitosis
• When cells reach a certain size, they
usually either stop growing or divide
• Cell cycle: stages through which a cell
passes from one cell division to the next
– Timing varies, usually 8 to 20 hours in actively
growing plant and animal cells
– Consists of two main phases, interphase and
M phase
© Cengage Learning 2015
Chromosomes Duplicate During
Interphase
• Most of a cell’s life is spent in interphase,
the time when no cell division is occurring
G1 S
(First gap (Synthesis
phase) phase)
G2
(Second
gap phase)
M PHASE
© Cengage Learning 2015 (Mitosis and cytokinesis)
ANIMATION: The Cell Cycle
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Three Phases of Interphase
• G1 phase
– Growth and normal metabolism; between the
end of mitosis and beginning of S phase
• S phase
– DNA replicates and histone proteins are
synthesized
• G2 phase
– Protein synthesis increases in preparation for
division
© Cengage Learning 2015
Interphase
Chromatin Nucleolus
Nucleus
Nuclear
envelope
Plasma
Centrioles membrane
© Cengage Learning 2015
3D Animation: Mitosis
M Phase
• Involves two main processes: mitosis and
cytokinesis
– Mitosis: nuclear division that produces two
nuclei containing chromosomes identical to
the parental nucleus
• Five stages: prophase, prometaphase, metaphase,
anaphase, and telophase
– Cytokinesis: division of the cell cytoplasm to
form two cells
© Cengage Learning 2015
During Prophase, Duplicated
Chromosomes Become Visible
• Begins with chromosome compaction
• Duplicated chromosomes consist of a pair
of identical sister chromatids
• Sister chromatids are held together at the
centromere by cohesins
• Attached to each centromere is a
kinetochore to which microtubules can
bind
© Cengage Learning 2015
Prophase (cont’d.)
• Each pole of the cell contains a
microtubule-organizing center from which
microtubules extend
– Animal cells have a pair of centrioles in the
middle of each microtubule-organizing center,
surrounded by pericentriolar material
– Microtubules radiate from pericentriolar
material toward the chromosomes, forming
the mitotic spindle
© Cengage Learning 2015
Prophase (cont’d.)
Sister chromatids
of duplicated
chromosome
Fragments
of nuclear
envelope
Developing
mitotic spindle
© Cengage Learning 2015
Prometaphase
• Begins when nuclear envelope breaks
down
• Spindle microtubules shorten or lengthen as
they contact chromosomes
– One of the kinetochores of a duplicated
chromosome “captures” a microtubule
– Chromosomes move toward cell’s midplane
– Cohesins dissociate from sister chromatid
arms, freeing them from one another
© Cengage Learning 2015
Prometaphase (cont’d.)
Spindle
microtubule
Kinetochore
© Cengage Learning 2015
Metaphase
• All the cell’s chromosomes align at the
cell’s metaphase plate
– Each chromatid is completely condensed
– The mitotic spindle has three types of
microtubules
– The remaining cohesin dissociate
• Chromosome abnormalities can be
detected using metaphase chromosomes
to construct a karyotype
© Cengage Learning 2015
Metaphase (cont’d.)
Spindle
Centriole
pair at
spindle
pole
Cell’s midplane
(metaphase plate)
© Cengage Learning 2015
The Mitotic Spindle
Metaphase plate (cell’s midplane)
Kinetochore microtubule
(chromosome spindle fiber)
Centrioles
Astral
microtubules
Pericentriolar
material
Polar (non-
kinetochore)
microtubule
Sister
chromatids
© Cengage Learning 2015
Anaphase
• Begins as sister chromatids separate into
separate chromosomes that move toward
opposite poles
• Kinetochore microtubules shorten at their
plus ends and pull chromosomes toward
the poles
• Ends when all chromosomes have
reached the poles
© Cengage Learning 2015
Anaphase (cont’d.)
Daughter chromosomes
© Cengage Learning 2015
Telophase
• Chromosomes arrive at the poles and
return to interphase-like conditions
• Chromosomes decondense by partially
uncoiling
• A new nuclear envelope forms around
each set of chromosomes
• Spindle microtubules disappear and
nucleoli reorganize
© Cengage Learning 2015
Telophase (cont’d.)
Cleavage furrow
Re-forming nuclear envelope
© Cengage Learning 2015
Cytokinesis
• Division of the cytoplasm to yield two
daughter cells
• Generally begins during telophase and
overlaps mitosis
• In animal or fungal cells:
– An actomyosin contractile ring attaches to
plasma membrane, causing constriction
– Cleavage furrow deepens and separates the
cytoplasm
© Cengage Learning 2015
Cytokinesis (cont’d.)
Cleavage
furrow
Actomyosin
contractile
ring
© Cengage Learning 2015
ANIMATION: Cytoplasmic Division
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Cytokinesis (cont’d.)
Cell plate
Nucleus forming
Vesicles
gather on Small vesicles Eventually one New cell walls
cell's fuse, forming large vesicle (from vesicle
midplane larger vesicles exists contents)
Plasma Cell Cell plate New plasma
membrane wall forming membranes
(from vesicle
membranes)
© Cengage Learning 2015
Daughter Cells of Mitosis
• Each daughter nucleus receives exactly
the same number and kinds of
chromosomes of parent cell
• Organelles are apportioned with cytoplasm
that each daughter cell receives during
cytokinesis
• Mitochondria and chloroplasts contain own
DNA and form by division of previously
existing mitochondria or plastids
© Cengage Learning 2015
Bacterial cell
Prokaryotes Divide Plasma 1
membrane Bacterial
by Binary Fission chromosome
Cell wall
Origin of
replication
Unreplicated
region of
Two copies chromosome 2
of bacterial
DNA
Z ring
New cell
walls
© Cengage Learning 2015 Two identical bacterial cells
Bacterial cell
Plasma 1
membrane Bacterial
chromosome
Cell wall
Origin of
replication
Unreplicated region
of chromosome
Two copies of
2
bacterial DNA
Z ring
New cell
walls
Stepped Art
Two identical bacterial cells Fig. 10-12, p. 224
ANIMATION: Prokaryotic Fission
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10.3 Regulation of the Cell Cycle
• Under optimal conditions, the length of the
eukaryotic cell cycle is constant for a given
cell type
• Cell-cycle checkpoints ensure all events of
a particular stage have been completed
before the next stage begins
• If a gene that encodes a molecule involved
in checkpoints is defective, cancer or other
serious diseases can result
© Cengage Learning 2015
Key Molecules in Cell Cycle Regulation
• Cyclin-dependent kinases (Cdks): protein
kinases that activate or inactivate other
proteins in by phosphorylating them
– Active only when they bind to cyclins to form a
cyclin–Cdk complex
• Eukaryotic cells form four major cyclin-Cdk
complexes:
– G1-Cdk, G1/S-Cdk, S-Cdk, M-Cdk
© Cengage Learning 2015
Cell Cycle Checkpoints (cont’d.)
1
G1 S
(First gap phase) (Synthesis phase)
G2
(Second gap
phase)
3
2
© Cengage Learning 2015
Other Key Molecules
• Anaphase-promoting complex (APC)
– Enzyme complex activated by M-Cdk
– Initiates anaphase by allowing degradation of
cohesins that hold sister chromatids together
• Hormones
– Plant hormones that promote mitosis in
normal growth and wound healing
– Certain steroids in animals can stimulate
growth and mitosis
© Cengage Learning 2015
Other Key Molecules (cont’d.)
• Protein growth factors
– Stimulate mitosis in some animal cells
– Examples: erythropoietin and epidermal
growth factor
• Certain drugs stop the cell cycle at a
specific checkpoint
– Some prevent DNA synthesis
– Some inhibit synthesis of proteins that control
the cycle or contribute to the mitotic spindle
© Cengage Learning 2015
10.4 Sexual Reproduction and Meiosis
• Asexual reproduction: a single parent
splits to produce a clone
– Rapid and efficient
– Favors organisms that are well adapted to
their environment
• Sexual reproduction: union of egg and
sperm to form a zygote
– Results in genetic variation among offspring
– Favors adaptation to changing environments
© Cengage Learning 2015
Homologous Chromosomes
• Paired chromosomes in somatic cells
– Similar in size, shape, and position of their
centromeres
– Carry information about the same genetic
traits, but not always the same information
– Human cells have 46 chromosomes in 23
homologous pairs
© Cengage Learning 2015
Diploid and Haploid Cells
• A set of chromosomes has one of each
kind of chromosome
• Diploid chromosome number: a cell or
nucleus contains two sets of
chromosomes
• Haploid number: a cell or nucleus has only
a single set of chromosomes
– In humans, the diploid chromosome number
is 46 and the haploid number is 23
© Cengage Learning 2015
Maintaining Chromosome Number
• Gametes are haploid
• When a sperm and egg fuse at
fertilization, each gamete (n) contributes
one set of chromosomes
– The diploid number is restored in the zygote
(2n)
• When the zygote divides by mitosis, all
somatic cells receive the same diploid set
of chromosomes
© Cengage Learning 2015
Polyploidy
• An individual whose cells have three or
more sets of chromosomes is polyploid
• Polyploidy is rare among animals but
common among plants
– As many as 80% of all flowering plants are
polyploid
– An important mechanism of plant evolution
© Cengage Learning 2015
Meiosis Produces Haploid Cells With
Unique Gene Combinations
• DNA in a diploid cell duplicates once, and
undergoes two divisions, yielding four
haploid nuclei
• Each haploid nucleus contains one
member of each homologous pair
• Each homologous chromosome pair is
shuffled, so resulting haploid cells each
have a unique combination of genes
© Cengage Learning 2015
Meiosis (cont’d.)
• Consists of two nuclear and cytoplasmic
divisions: meiosis I and meiosis II
– Each includes prophase, metaphase,
anaphase, and telophase stages
– Meiosis I: partner homologous chromosomes
pair, then separate and move into different
nuclei
– Meiosis II: sister chromatids separate from
each other and are distributed to two different
nuclei
© Cengage Learning 2015
Interphase
One 2n cell with four
unduplicated
chromosomes (2
homologous pairs)
Replication of DNA
Four chromosomes, Homologous pair of
each with two sister duplicated chromosomes
chromatids
Meiosis I: Homologous chromosomes
pair and then separate
Sister chromatids
Meiosis II:
Sister chromatids separate
Prophase I Includes Synapsis and
Crossing-over
• Each duplicated chromosome consists of
two chromatids, linked by cohesins
• Homologous chromosomes align side by
side, which results in association of four
chromatids
• A synaptonemal complex forms along the
length of the synapsed homologues
• Enzymes break and rejoin DNA molecules,
allowing crossing-over
© Cengage Learning 2015
Prophase I (cont’d.)
INTERPHASE MEIOSIS I
Prophase I
Nucleolus Homologous
chromosomes
Nuclear envelope
Chromatin
Developing
Centrioles
meiotic spindle
Interphase preceding Homologous chromosomes
meiosis; DNA replicates. synapse and exchange segments
by crossing over; nuclear envelope
breaks down.
© Cengage Learning 2015
During Meiosis I, Homologous
Chromosomes Separate
Metaphase I Anaphase I Telophase I
Microtubule
attached to Cleavage furrow
kinetochore
Sister Separation of
chromatids homologous
chromosomes
Tetrads line up on Homologous chromosomes One of each pair
cell's midplane. separate and move to of homologous
Tetrads held together opposite poles. Note that chromosomes is
at chiasmata (sites of sister chromatids remain at each pole.
prior crossing-over). attached at their Cytokinesis occurs.
© Cengage Learning 2015 centromeres.
Meiosis II
MEIOSIS II Metaphase II
Prophase II
Chromosomes condense again Chromosomes line up along
following brief period of interkinesis. cell's midplane.
DNA does not replicate again.
© Cengage Learning 2015
Meiosis II (cont’d.)
Anaphase II Telophase II Four haploid cells
Daughter
chromosomes
Sister chromatids Nuclei form at Four gametes
separate, and opposite poles of (animal) or four
chromosomes move each cell. Cytokinesis spores (plant) are
to opposite poles. occurs. produced.
© Cengage Learning 2015
3D Animation: Meiosis
Meiosis Produces Genetic Variation
• Two successive divisions of meiosis yield
four haploid nuclei, each with a different
combination of genes
– Two sources:
• DNA segments are exchanged between maternal
and paternal homologues during crossing-over
• Maternal and paternal chromosomes of
homologous pairs separate independently at
anaphase I
© Cengage Learning 2015
Mitosis and Meiosis Lead to
Contrasting Outcomes
• Mitosis is a single nuclear division which
produces two daughter cells that are
genetically identical to each other and to
the original cell
• In meiosis, a diploid cell undergoes
genetic rearrangement and two
successive nuclear divisions, resulting in
formation of four genetically different,
haploid daughter cells
© Cengage Learning 2015
10.5 Sexual Life Cycles
• In animals, somatic cells are diploid and
multiply by mitosis
– Only haploid cells produced are gametes
– Gametes develop when germ line cells
undergo meiosis
• Gametogenesis: formation of gametes
• Spermatogenesis: forms four haploid sperm cells
for each cell that enters meiosis
• Oogenesis: forms one egg cell for every cell that
enters meiosis, plus polar bodies
© Cengage Learning 2015
Animal Life Cycles
Gametes (n)
Meiosis Fertilization
Zygote (2n)
Mitosis
Multicellular
diploid
organism
(a) Animals (2n)
© Cengage Learning 2015
Life Cycles in Most Fungi and Many
Protists
Unicellular or
multicellular
haploid organism
(n)
Mitosis Mitosis
Gametes (n)
Meiosis Fertilization
Zygote (2n)
(b) Most fungi and many protists
© Cengage Learning 2015
Life Cycles in Plants
• Complicated life cycles with alternation of
generations
– Cells of a multicellular 2n sporophyte undergo
meiosis to form haploid spores, each of which
may divide mitotically to produce a
multicellular haploid gametophyte
– Gametophytes produce female and male
gametes by mitosis; egg and sperm fuse to
form 2n zygote that divides mitotically to form
a multicellular sporophyte
© Cengage Learning 2015
Life Cycles in Plants (cont’d.)
Gametophyte (n)
(multicellular
haploid organism)
Mitosis Mitosis
Spores (n) Gametes (n)
Meiosis Fertilization
Zygote (2n)
Mitosis
Sporophyte (2n)
(multicellular
diploid organism)
© Cengage Learning 2015 (c) Plants, some algae, and some fungi
ANIMATION: Generalized Life Cycles
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VIDEO: The Discovery of How Cells Divide
VIDEO: Meiosis and Genetic Hotspots
ANIMATION: Mitosis and Meiosis Compared