CHAPTER 2

MITOSIS & MEIOSIS

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Chapter 2 Outline – Learning

Outcomes
1. Cell Structure Is Closely Tied to Genetic Function
2. Chromosomes Exist in Homologous Pairs in Diploid Organisms
3. Mitosis Partitions Chromosomes into Dividing Cells
4. Meiosis Creates Haploid Gametes and Spores and Enhances Genetic
Variation in Species
5. The Development of Gametes Varies in Spermatogenesis Compared to
Oogenesis
6. Meiosis Is Critical to the Successful Sexual Reproduction of All Diploid
Organisms
7. Electron Microscopy Has Revealed the Physical Nature of Mitotic and
Meiotic Chromosomes

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  Living organisms have genetic material
 Composed of nucleic acid DNA
 Organized into chromosomes
 Does not include viruses
 In eukaryotes, transmission of genetic material via chromosomes involves
mitosis and meiosis
 Meiosis leads to production of sex cells:
 Gametes or spores
 Mitosis leads to production of two cells
 Each with same number of chromosomes as parent cell
 During nondivisional phases, chromosomes uncoil into a diffuse network within the
nucleus: chromatin

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I- Cell Structure is Closely Tied to

Genetic Function

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 There are two main types of cells
 Prokaryotic (bacteria, archaea)
 Eukaryotic (protists, plants, fungi, animals)

All cells share some common features

 Plasma membrane
 DNA
 Ribosomes

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 I.1- Plasma Membrane

 Surrounds all cells

 Delimits cell from external environment
 Plants
 Have plasma membrane
 Have a cell wall
 Composed mainly of cellulose, a polysaccharide
 Bacterial cells have peptidoglycan in their cell walls

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I.2- Glycocalyx

 Glycocalyx or Cell Coat

 Covering on plasma membrane of animal cells
 Made of glycoproteins and polysaccharides
 Function: biochemical identity at cell surface
 Receptor Molecules are also found on the surface of cells
 Recognition sites that transfer specific chemical signals across the cell
membrane into the cell

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 I.3- Nucleus and Nucleoid

 Nucleus
 Found in eukaryotes
 Membrane bound
 Houses genetic material, DNA
 Nucleolus: where ribosomal RNA (rRNA) is synthesized
 Nucleolus organizer region (NOR): DNA that encodes rRNA
 Nucleoid
 Found in prokaryotes
 Not membrane bound

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I.4- Cytoplasm

 Cytoplasm includes:
 Extra nuclear cellular organelles
 Cytosol: Colloidal material surrounding organelles
 Cytoskeleton made of extensive system of tubules and filaments
 Microtubules: Made up of the protein tubulin
 Microfilaments: Derived from the protein actin
 Intermediate Filaments (eg. keratins)

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 I. The Endoplasmic Reticulum

 ER: An organelle
 Compartmentalizes cytoplasm
 Increases surface area for biochemical synthesis
 SER: Smooth ER appears smooth in places
 Site of fatty acid and phospholipid synthesis
 RER: Rough ER is studded with ribosomes
 Site of protein synthesis

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I.6- Cytoplasmic Structures- Mitochondria

and Chloroplast
 Mitochondria: Animal and plant cells
 Site of oxidative phases of cell respiration, which generate ATP
 Chloroplasts: Plants, algae, and protozoans
 Site of Photosynthesis

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 I.7- Centrioles

 Found in centrosome of animal cells

 Organize spindle fibers for movement of chromosomes during meiosis and mitosis
 Made up of microtubules

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II- Chromosomes Exist in Homologous

Pairs in Diploid Organisms

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 II.1- Homologous Chromosomes

 Chromosome = DNA + packing proteins

 Chromosomes exist in homologous pairs in diploid organisms
 Homologous chromosomes
 Are similar but not identical
 Carry genes for the same inherited characteristics
 May carry different versions of the same gene
 Called alleles

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telomere
p arm

centromere

q arm

telomere

Chromosome at the beginning of mitosis

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 II.2- Centromere

 Centromeres are constricted regions on chromosomes

 Location of centromere establishes appearance of chromosome
 Metacentric
 Submetacentric
 Acrocentric
 Telocentric
 Sister chromatids with centromere
 Long arm = q arm
 Short arm = p arm

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 II.3- Ploidy

 Ploidy = number of chromosome sets in a cell.

 Haploid = n chromosomes
 Diploid = 2n chromosomes organized in n pairs
 Polyploid (3n, 4n, 6n…)
 Aneuploid = abnormal number of chromosomes

 Somatic cells (body cells) have homologous pairs of chromosomes

 Humans: 46 chromosomes (23 homologous pairs)
 Represents diploid number, 2n
 Exceptions:
 Many bacteria and viruses have one chromosome

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II.3- Ploidy
 Karyotype:
 Illustrates the physical appearance of the different pairs of homologous
chromosomes
 The human mitotic chromosomes have been photographed, cut out of the print,
and matched up

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 II.3- Ploidy

 Meiosis converts the diploid number (2n) of chromosomes to the haploid

number (n)
 Genome: Genetic information in haploid set

 Homologous chromosomes contain identical gene sites along their lengths

 Each site is called a locus (p l. loci)
 Biparental inheritance: inheritance from two parents
 Diploid organisms contain two copies of each gene
 Alleles: alternative forms of the same gene can exist

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 II.4- Gonosomes

 Sex-determining chromosomes
 Not homologous
 Behave as homologs in meiosis
 Example:
 X and Y chromosome in humans

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III- Mitosis Partitions Chromosomes

into Dividing Cells

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  Critical process for all Eukaryotes
 Reproduction of unicellular organisms
 Development of multicellular organisms from zygote (single-celled fertilized egg)

1. Karyokinesis: genetic material partitioned to daughter cells during nuclear division

2. Cytokinesis: cytoplasmic division follows

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III.1- Interphase and Cell Cycle

 Cell cycle
 Composed of interphase and mitosis

G1 = Gap 1
S = Synthesis
G2 = Gap 2

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 III.2- Interphase

 Interphase includes
 S phase: DNA is synthesized
 Two gap phases (G1 and G2)
 G0: Point in G1 phase where cells are nondividing, but a metabolically
active state
 Interphase: characterized by the absence of visible chromosomes

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III.2- Interphase

 Cell cycle length varies:

 Mitosis <1h
 S and G2 are almost constant in a species
 G1 extremely variable
 Cancer cells never undergo G0
 Some cells never exit G0 (neurons)
 Most cells are temporarily in G0 and return to cell cycle when required

 Mitosis: prophase, prometaphase, metaphase, anaphase, telophase

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 III.3- Mitosis

Prophase
1. In animal cells: Centrioles divide, move, establish poles
 Plant cells form their spindle fibers w/o centrioles
2. Nuclear envelope breaks down
3. Chromosomes condense and become visible
 Sister chromatids: Two parts of each chromosome
 Connected at the centromere
 Held together by multi-subunit protein complex called cohesin

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III.3- Mitosis

Prometaphase
 Period of chromosome movement to equatorial
plane of cell
 Equatorial plane referred to as metaphase plate
 Centrioles reach poles
 Spindle fibers form

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 III.3- Mitosis

Metaphase
 Chromosome configuration following migration
 Centromeres align on metaphase plate

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III.3- Mitosis

Metaphase
 Kinetochore
 Proteins associated with centromere
 Spindle fibers bind to kinetochore; chromosomes migrate
 Kinetochore microtubules: have one end near the centrosome region and the
other end anchored to the kinetochore

 Cohesin
 Protein complex that holds sister chromatids together
 Separase
 Enzyme that degrades cohesin
 Shugoshin
 Protein that protects cohesin from being degraded by separase
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III.3- Mitosis

Anaphase = Disjunction
 Sister chromatids separate now called daughter chromosomes
 Migrate to opposite poles by shortening of spindle fibers
 Motor proteins or molecular motors use ATP

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 III.3- Mitosis

Telophase
 Cytokinesis
 Uncoiling of the chromosomes
 Re-formation of the nuclear envelope
 Spindle fibers disappear
 Nuclear envelope reforms

Cytokinesis
 Two new cells are produced
 Plant cells: a cell plate is synthesized and laid down across the metaphase plate
 Becomes the middle lamella
 Animal cells: a constriction of the cytoplasm or cell furrow
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III.3- Mitosis

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 III.3- Mitosis

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III.3- Mitosis

Summary of mitosis
 Partitions chromosomes into dividing cells
 Produces daughter cells with a full diploid complement of chromosomes

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 III.3- Mitosis

Summary of mitosis
 Partitions chromosomes into dividing cells
 Produces daughter cells with a full diploid complement of chromosomes

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III.4- Cell Cycle Regulation and Checkpoints

 Many mutations are now known that exert an effect on the cell cycle
 Cell division cycle (cdc) mutations: enzymes called kinases
 Can add phosphates to other proteins to regulate the cell cycle
 Are “master control” molecules functioning in conjunction with cyclin proteins
 The cell cycle checkpoints: monitor mitosis for errors

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 III.4- Cell Cycle Regulation and Checkpoints

 G1/S checkpoint
 Monitors size cell has achieved
 Evaluates condition of DNA
 G2/M checkpoint
 Monitors if DNA replication is incomplete
 Monitors damaged DNA
 M checkpoint
 Monitors successful formation of spindle fiber system and attachment to kinetochores

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IV- Meiosis Creates Haploid Gametes

and Spores and Enhances Genetic
Variation in Species

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  Mitosis maintains diploidy and amount of genetic material
 Meiosis
 Reduces amount of genetic material by half
 Produces haploid gametes or spores, each containing one member of a
homologous pair of chromosomes
 Crossing Over
 Meiotic event
 Genetic exchange between members of homologous pairs of chromosomes

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IV.1- Overview of Meiosis

 Meiosis I: Reductional division

 Meiosis II: Equational division

 DNA synthesis
 Occurs during interphase, before the beginning of meiosis I
 Does not occur again before meiosis II

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 IV.1- Overview of Meiosis

 Meiosis I and II
 Each has prophase, metaphase, anaphase, and telophase stages

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 IV.2- Meiosis I – Prophase I

 3 main events:
 Chromatin condenses into chromosomes
 Pairing of homologous chromosomes
 Crossing-over between paired chromosomes
 Five substages, each with specific events:
 Leptonema, Zygonema, Pachynema, Diplonema, Diakinesis
 End of prophase I:
 Centromeres are present on the equatorial plate for
Metaphase

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IV.2- Meiosis I – Prophase I

1. Leptonema
 Chromatin starts to condense
 Chromosomes appear as long, single threads, unassociated
with each other
 Darker regions = chromomeres

2. Zygonema
 Ongoing condensation and thickening of chromosomes
 Pairing of homologous chromosomes = Synapsis
 Each pair of homologous chromosomes is known as bivalent
 n bivalents

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 IV.2- Meiosis I – Prophase I

3. Pachynema
 Each bivalent becomes shorter, thicker, and splits into two
sister chromatids called tetrads

 Crossing over
 Exchange of genetic material between nonsister chromatids
 In areas called chiasmata

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IV.2- Meiosis I – Prophase I

4. Diplonema
 Pulling apart of each pair of sister chromatids except for
chiasmata

5. Diakinesis
 Nucleus and nuclear envelope break down
 Centromeres attach to spindle fibers

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 IV.3- Meiosis I: Metaphase I

 Chromosomes at maximum shortness

 Terminal chiasmata holding nonsister chromatids together
 Binding to spindle fibers moves chromatids to metaphase plate

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IV.3- Meiosis I: Anaphase I

 Cohesin is degraded between sister chromatids

 Homologous chromosomes separate
 Nondisjunction may occur

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 IV.3- Meiosis I: Telophase I

 In many organisms:
 Nuclear membrane forms
 Nucleus enters interphase (w/o DNA replication)
 In other organisms: directly from anaphase I to meiosis II

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IV.3- Meiosis I: Summary of Metaphase I,

Anaphase I and Telophase I

 Homologous chromosomes separate and move toward the poles

 Sister chromatids remain attached at centromeres
 Duplicated chromosomes reach poles. Each pole now has haploid set of
chromosomes
 Cytokinesis: Two haploid daughter cells are formed
 Nuclear envelope forms around chromosomes in some species

 Meiosis significantly increases the level of genetic variation due to crossing over

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IV.4- Meiosis II – Second Meiotic Division

 Second meiotic division – equational (similar to mitosis)

 During Prophase II: each dyad is composed of one pair of sister chromatids
attached by the common centromeric region
 During Metaphase II: the centromeres are positioned on the equatorial plate
 During Anaphase II: sister chromatids are separated to opposite poles
 Each haploid daughter cell from meiosis II has one member of each pair of homologous
chromosomes
 During Telophase II: one member of each pair of homologous chromosomes present
at each pole
 Each chromosome is now a monad
 Cytokinesis results in 4 haploid cells

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V- The Development of Gametes

Varies in Spermatogenesis Compared
to Oogenesis

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  Similar meiotic events in all cells participating in gametogenesis in most animal
species
 The development of gametes varies between spermatogenesis and oogenesis

 Male gametes: Produced by spermatogenesis in the testes

 Female gametes: produced by oogenesis in the ovary

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 V.1- Spermatogenesis

 In testes
 May be continuous or may occur periodically in mature male animals depending
on reproductive cycles (year-round or seasonal).
 Growth of undifferentiated diploid germ cell = spermatogonium
 Primary spermatocyte undergoes meiosis I
 Produces two secondary spermatocytes
 Spermatocytes undergo meiosis II
 Produces a total of four haploid spermatids (equal amounts of cytoplasm)
 Spermatids undergo spermiogenesis
 Series of developmental changes
 Become highly specialized motile spermatozoa or sperm

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V.2- Oogenesis

 Formation of ova in the ovaries

 Four daughter cells do not receive equal cytoplasm
 Only one daughter cell receives cytoplasm (+polar body)
 Undergoes meiosis I and II
 Develops into ovum

 Discontinuous in many animal species, including humans.

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 oocyte II

oocyte I First polar

body

Prophase I Metaphase I Anaphase I Telophase I Cytokinesis

Embryonic life Puberty

Arrest

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ootid

oocyte II

Second
polar body
Prophase II Metaphase II Anaphase II Telophase II Cytokinesis

Completion after
Arrest-
ovulation fertilization

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 VI- Meiosis Is Critical to the Successful
Sexual Reproduction of All Diploid
Organisms

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VI.1- Importance of meiosis

 Meiosis
 Mechanism by which diploid amount of genetic information is reduced to haploid
 Fertilization restores diploidy.

 Great amount of genetic reshuffling:

 Paternal or maternal chromosome of each pair in the gamete
 Crossing-over = reshuffling of alleles between homologs

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 VI.2- In Fungi and Plants

 Plant and fungi life cycles

 Fungi: Haploid vegetative cells arise via meiosis—proliferate via mitotic cell division
 Plants: Life cycle alternates between diploid sporophyte stage and haploid gametophyte
stage

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 VII- Electron Microscopy Has Revealed
the Physical Nature of Mitotic and
Meiotic Chromosomes

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VII.1- Chromosomes

 Visible only during mitosis and meiosis

 Chromatin fibers that make up chromosomes coil and condense
 EM of mitotic chromosomes in varying states of coiling led to the folded-fiber model

(a) chromatin fibers characteristic of the interphase nucleus (TEM) (b) metaphase chromosomes that are
derived from chromatin during mitosis (SEM) (c) diagram of a mitotic chromosome, showing how
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 VII.2- DNA folding

Chromosomes are composed of DNA

tightly-wound around histones.
© 2013 Nature Education

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