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MEIOSIS
THE PROCESS OF REDUCTION DIVISION
Submitted by,
A.T Milin Sera
Roll no : 1
1st M.sc. Botany
ST. Teresa’s College, Ekm
Submitted to,
Dr. Arya P Mohan
Assistant Professor
ST. Teresa’s College, Ekm
CONTENTS
• INTRODUCTION
• MEIOSIS
• PHASES OF MEIOSIS
• MEIOSIS VS MITOSIS
Meiosis...
INTRODUCTION
• The cells of a particular species have a constant number of chromosomes.
• If the gametes had the same number of chromosomes as the somatic cells, then
the zygote would have twice the diploid number of chromosomes.
• This number would go on doubling with each generation.
• However, the chromosome number remains constant from generation to
generation.
• This is because of meiotic division which reduces the chromosome number to
half, and counteracts the effect of fertilization.
• Thus fertilization and meiosis are compensating events.
Meiosis...
1
MEIOSIS
• Extracellular signals induce a transcriptional program that produces
meiosis-specific cell cycle.
• Such an extracellular signals inducing entry into the meiotic divisions
in mammals is retinoic acid, which is a steroid hormone.
• Meiosis consists of two cell divisions.
• The two divisions are known as the first meiotic division and the
second meiotic division.
• The stages of first meiotic division are prophase I, prometaphase I,
metaphase I, anaphase I and telophase I.
• The second meiotic division consists of prophase II, metaphase II,
anaphase II and telophase II.
Meiosis...
2
PHASES OF MEIOSIS
• Before entering meiosis I, a cell first go through interphase.
• The interphase preceding meiosis is important because
replication of DNA takes place during this stage.
• Replication is confined to the synthesis (S) phase of
interphase.
• It is preceded by a post-mitotic gap phase (G1) and followed
by a pre-mitotic gap phase (G2).
• During these two phases there is no replication of DNA.
PRE-MEIOTIC INTERPHASE
3
Meiosis...
FIRST MEIOTIC DIVISION (MEIOSIS-1)
• Cell division that reduces the chromosome number by one-half.
• It separate homologous chromosomes and produce two cells with
haploid chromosome number (n).
• For that reason it is known as Reductional Division.
• Four phases: Prophase I, Metaphase I, Anaphase I,Telophase I
PROPHASE -1
• Longest and most complex phase.
• It consists of 5 substages : Leptotene, Zygotene, Pachytene,
Diplotene, Diakinesis.
• In G2 and prophase of meiosis I, the two replicated chromatids of each
chromosome are associated with each other by cohesin complexes.
4
Meiosis...
LEPTOTENE
• Chromsomes become very thin
• They are not distinct, cannot be easily seen.
PROLEPTOTENE
• During this stage the chromosomes become more distinct.
• Chromosomes appear as slender threads bearing a series of
granule- like structures called chromomeres.
• Under electron microscope the leptotene chromosomes
have an axial filament.
• To this filament, chromatin fibres are attached as a series of
lateral loops.
• Cytoplasm has many polyribosomes, but few ER.
• A pre-meiotic pairing may takes place during leptotene.
5
Meiosis...
• Leptotene chromosomes may be irregularly arranged, or may be polarized
towards the centrioles forming a ‘bouquet’.
• In plant cells the chromosomes may sometimes form a tangle of threads, called
the synizetic knot, on one side of the nucleus.
• During leptotene the cytoplasm has many polyribosomes, but endoplasmic
vesicles are few.
• A pre-meiotic pairing may takes place during leptotene.
• Homologous chromosomes may be aligned together, but no synaptonemal
complexes are formed.
6
Meiosis...
ZYGOTENE
• During zygotene the chromosomes become shorter and
thicker.
• The homologous chromosome starts pairing together.
• These homologous chromosome come to lie side by side in
pairs & this pairing is called synapsis.
• The complex formed by a pair of synapsed chromosome is
called bivalent.
• Pairing may begin at the centromere and proceed towards the
ends (procentric pairing).
• They may begin at the ends and proceed towards the
centromere (proterminal pairing).
7
Meiosis...
• Sometimes it may occur simultaneously at all chromomeres (intermediate
condition).
• In many plant and animal species a kind of pseudosynapsis between non-
homologous regions also occurs.
• When synapsing occurs, the chromosomes do not lie side by side but rather on
top of each other with a protein called the synaptonemal complex holding them
together.
8
Meiosis...
SYNAPTONEMAL COMPLEX
• The synaptonemal complex is a proteinaceous
structure that is formed between paired
homologous chromosomes.
• It is a tripartite ribbon like structure.
• It consists of a dense central element, with a dense
lateral element on either side.
• The lateral elements are attached at both ends to
the nuclear membrane.
• The lateral elements are connected to the central
element by a series of lateral loops which form
transverse units or LC fibres.
9
Meiosis...
• Three specific components of the synaptonemal complex have been characterized:
SC protein-1 (SYCP1), SC protein-2 (SYCP2), and SC protein-3 (SYCP3).
• Lateral elements are mainly formed by SYCP3 and secondarily by SYCP2.
• Central element and transverse filaments are composed by the protein SYCP1.
• The axial filaments of leptotene chromosomes later become transformed into the
lateral elements of the SCs.
• The SCs first appear during zygotene.
• They are completed by the beginning of pachytene.
• By diplotene the SCs have mostly disappeared, but remnants may be seen even in
diakinesis.
• They are believed to be involved in chiasmata formation and crossing over.
• It stabilise the two homologous chromosomes till the crossing over is completed.
Meiosis...
10
PACHYTENE
• The bivalents appear as thick threads due to condensation of
chromosomes.
• Two chromatids belonging to different homologues undergo
one or more transverse breaks at the same level.
• The breaks never occur between two sister chromatids.
• The break is followed by interchange and fusion of broken
ends between two homologous chromosomes.
• This is called crossing over.
• The site where crossing over occurs forms a recombination
nodule.
• An enzyme called recombinase is involved during this phase.
11
Meiosis...
DIPLOTENE
• Separation of paired homologues chromosomes begin.
• The separating chromosomes are held together at one or more
points called chiasmata.
• A chiasma formed at the ends of chromosomes is called a
terminal chiasma.
• Chiasmata formed along the lengths of chromosomes are called
interstitial chiasmata.
• Terminalization of chiasmata takes place.
• The synaptonemal complexes mostly disappear during diplotene.
• Cytoplasm shows an increase in the number of endoplasmic
vesicles.
Meiosis...
12
DIAKINESIS
• The chromosomes become more contracted.
• The bivalents are more evenly distributed in the nucleus
and migrate towards the periphery.
• The homologues remain in contact with each other by
their terminal chiasmata.
• The nucleolus is detached from the chromosome, or
disappears.
• Remnants of the synaptonemal complexes may still be
seen.
13
Meiosis...
METAPHASE-1
• The chromosomes now become arranged on the
equator of the cell.
• The spindle is formed.
• Spindle fibres become attached to the centromeres of
the two homologous chromosomes.
• The two centromeres of each bivalent lie on opposite
sides of the equatorial plate.
• The nuclear membrane disappears.
• The chromosomes reach their maximum contraction.
• Spindle formation begins.
PROMETAPHASE
1
Meiosis...
14
ANAPHASE-1
• Separation of the two homologous chromosomes of each
bivalent begins.
• One chromosome from each bivalent begins to migrate to one
pole, the other migrates to the opposite pole.
• The sister chromatids are said to be co-oriented.
• The kinetochores of each bivalent attach to spindle microtubules
from opposite spindle poles.
• They are said to be bi-oriented.
• During Anaphase I original chromosomes separate, so reduction
in the number of chromosomes from 2n to n number.
15
Meiosis...
TELOPHASE-1
• The homologous chromosome complete their migration to the
two poles because of shortening of spindles.
• When the chromosomes reach the poles they persist for some
time in the condensed state.
• Later they undergo despiralization and become elongated.
• The nuclear membrane is reformed, but the nucleoli do not
reappear.
• Unlike in mitosis the spindle fibres do not disappear
completely.
16
Meiosis...
CYTOKINESIS
• Cytokinesis involves the formation of a cleavage furrow, resulting in the
pocketing of the cell into two cells.
• In most plant cells daughter cells are formed by the laying down of cell plate
between the two groups of chromosomes.
• In other plant cells cytokinesis does not take place until both meiotic divisions are
completed.
• At the end of Telophase I and Cytokinesis, two daughter cells are produced.
• Each daughter cell with one half of the number of chromosomes of the original
parent cell.
Meiosis...
17
18
Meiosis...
MEIOSIS I STAGES MAIN EVENTS
Prophase 1 : Sub-Stages :-
Leptotene Chromosomes appear as slender threads
& Bouquet Formation
Zygotene Synapsis & synaptonemal complex
formation
Pachytene Chromosomes appear as thick threads &
Crossing over
Diplotene Chiasma formation
Diakinesis Chromosomes get thick and contracted
Metaphase 1 Chromosomes arranged in equator region
& Spindle formation
Anaphase 1 Separation of homologous chromosomes
& Beginning of chromosomes migration to
opposite poles
Telophase 1 Migration of homologous chromosomes
complete & Shortening of spindle fibres
INTERKINESIS OR INTRAMEIOTIC
INTERPHASE
• Interkinesis is a metabolic phase between telophase-1 & prophase-2.
• During this phase chromosomes are elongated but do not form chromatin
fibres.
• This differs from interphase I and interphase of mitosis in that there is no
duplication of DNA.
• The RNA and protein required during meiosis-2 are syntesised during this
phase.
19
Meiosis...
MEIOSIS -2
• During Meiosis II, two sister chromotids of each chromosome separate and
migrate to the opposite pole.
• As a result, the number of chromosomes in each of the two haploid nuclei
remains the same (i.e haploid), at the end of this division.
• Thus the second division of meiosis is often referred to as equational division.
• Sometimes, it is called as ‘Meiotic Mitosis’.
• The second meiotic division is also divided into four stages : Prophase II,
Metaphase II, Anaphase II, Telophase II.
• The 4 stages of meiosis – II is same as mitosis.
• It differs from mitosis in that DNA does not duplicate, while the centromeres do
so.
Meiosis...
20
21
Meiosis...
MEIOSIS II STAGES MAIN EVENTS
Prophase 2 Chromatids with widely separated
arms & Spindle formation and
nuclear membrane disappearance
Metaphase 2 Chromosomes oriented at
equatorial plate & Spindle fibre
attachment as same as in mitosis
Anaphase 2 Centromeres divide & Two
chromatids of each chromosome
separate to opposite poles
Telophase 2 Reconstruction of nuclei. Nucleus,
centriole and chromosomes return
to interphasic condition & Nucleus
contains haploid no. of
chromosomes
Cytokinesis Dyad divides into two parts. One
parent cell produce four haploid
daughter cells, together known as
tetrad
REMOVAL OF COHESION
• To facilitate two consecutive chromosome segregation phases, cohesins have to be
lost from chromosome.
• During meiosis, cohesins are lost from chromosome arms by the end of meiosis I,
but a pool of cohesins around kinetochores is protected from removal.
• This pool of cohesins persists throughout meiosis I but is removed at the onset of
anaphase II.
• The mechanisms that remove cohesins during meiosis are the same as during
mitosis.
• Securin degradation releases separase, which then cleaves the cohesins.
• This allows the recombined maternal and paternal chromosomes to separate, but
each pair of chromatids remains associated at the centromere.
• During metaphase II, sister chromatids align on the metaphase II spindle and
separase is activated yet again, cleaving the residual cohesin around centromeres.
22
Meiosis...
Rec8
• A specialized cohesin subunit, Rec8, is necessary for the stepwise loss of cohesins
from chromosomes during meiosis.
• Rec8 is homologous to Scc1, the cohesin subunit that closes the cohesin ring in
the cohesin complex of mitotic cells.
• During early anaphase of meiosis I, Rec8 is lost from chromosome arms but is
retained at centromeres.
• During early anaphase of meiosis II, centromeric Rec8 is cleaved by separase, so
the sister chromatids can segregate.
• The mechanism that protects Rec8 from cleavage at centromeres during meiosis I
is similar to the mechanism that protects Scc1 at centromeres during mitosis.
Meiosis...
23
PP2A & SHUGOSHIN
• Cohesion at the centromeres is maintained because a specific isoform of protein
phosphatase 2A (PP2A).
• PP2A is localized to centromeric chromatin by members of a family of proteins
known as the Mei-S332/Shugoshin.
• PP2A keeps cohesin in a hypophosphorylated state that does not dissociate.
• During metaphase II, Mei-S332/Sgo1 dissociates from chromosomes.
• When the last kinetochore is properly associated with spindle microtubules,
Cdc20 is depressed and associates with the APC/C.
• This causes ubiquitinylation of securin.
• This releases separase activity, which cleaves Rec8, eliminating cohesion at the
centromere and allowing chromatid separation in anaphase.
24
Meiosis...
• Cohesin removal differs for meiosis I because when Rec8
replaces Scc1 in the cohesin complex.
• The meiotic cohesin complex can only be removed from
chromatin via the action of separase.
• Rec8 also differs from Scc1 in that it must be phosphorylated by
several protein kinases to be cleaved by separase.
• During meiosis I, the centromere-specific isoform of PP2A
targeted to centromeric chromatin by Mei-5332/Shugoshin
prevents this phosphorylation.
• The PP2A targeting factor and PP2A then dissociate from
chromosomes by metaphase II, allowing separase cleavage of
Rec8.
Meiosis...
25
• In mitosis and meiosis ll, sister kinetochores attach to spindle microtubules
emerging from opposite spindle poles.
• These kinetochores are said to be bi-oriented.
• This is essential for segregation of sister chromatids to different daughter cells.
• In contrast, at meiosis I metaphase, sister kinetochores attach to spindle
microtubules emerging from the same spindle pole.
• These sister kinetochores are said to be co-oriented.
• Attachment of sister kinetochores to the proper microtubules in meiosis I and II is
critical for correct meiotic segregation of chromosomes.
ORIENTATION OF KINETOCHORES
26
Meiosis...
• Monopolin complex is a protein complex that links with sister kinetochores
during meiosis I for their attachment to microtubules emerging from the same
spindle pole.
• In organisms where kinetochores attach to multiple microtubules, Rec8-
containing cohesins are essential for sister kinetochore co-orientation.
• These meiosis-specific cohesins impose a rigid kinetochore structure.
• This restricts the movement of sister kinetochores and thereby favoring
attachment to microtubules from the same spindle pole.
• Like during mitosis and meiosis II, correct attachment of meiosis I chromosomes
is mediated by a tension-based mechanism.
• During meiotic metaphase I, kinetochore-associated microtubules are also under
tension.
Meiosis...
27
• Because chiasmata generated by recombination between
homologous chromosomes and the cohesins distal to the
chiasmata prevent them from being pulled to the poles.
• Kinetochore-microtubule attachments are unstable in the
absence of tension.
• When kinetochores attach to the wrong spindle fibers
release the incorrect microtubules.
• This enables them to bind microtubules again until
attachments are made that generate tension.
• As in mitosis, once tension is generated, microtubule
attachment to the kinetochores is stabilized.
28
Meiosis...
MEIOSIS VS MITOSIS
29
Meiosis...
REFERENCE
• Burgess, J. (1985). Introduction to plant cell development. CUP Archive.
• Lodish, H. (2008). Molecular Cell Biology. New York: W.H. Freeman and Company.
• Pawar, C.B. (1983). Cell Biology. Himalaya Publishing House.
• https://www.slideshare.net/KAUSHIKKUMARPANIGRAH/presentation-on-meiosis
• https://www.google.com/amp/s/slideplayer.com/amp/14961858/
• https://www.khanacademy.org/science/ap-biology/heredity/meiosis-and-
genetic-diversity/a/phases-of-meiosis
30
Meiosis...
THANK YOU

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Meiosis : introduction and phases of meiosis.

  • 1. MEIOSIS THE PROCESS OF REDUCTION DIVISION Submitted by, A.T Milin Sera Roll no : 1 1st M.sc. Botany ST. Teresa’s College, Ekm Submitted to, Dr. Arya P Mohan Assistant Professor ST. Teresa’s College, Ekm
  • 2. CONTENTS • INTRODUCTION • MEIOSIS • PHASES OF MEIOSIS • MEIOSIS VS MITOSIS Meiosis...
  • 3. INTRODUCTION • The cells of a particular species have a constant number of chromosomes. • If the gametes had the same number of chromosomes as the somatic cells, then the zygote would have twice the diploid number of chromosomes. • This number would go on doubling with each generation. • However, the chromosome number remains constant from generation to generation. • This is because of meiotic division which reduces the chromosome number to half, and counteracts the effect of fertilization. • Thus fertilization and meiosis are compensating events. Meiosis... 1
  • 4. MEIOSIS • Extracellular signals induce a transcriptional program that produces meiosis-specific cell cycle. • Such an extracellular signals inducing entry into the meiotic divisions in mammals is retinoic acid, which is a steroid hormone. • Meiosis consists of two cell divisions. • The two divisions are known as the first meiotic division and the second meiotic division. • The stages of first meiotic division are prophase I, prometaphase I, metaphase I, anaphase I and telophase I. • The second meiotic division consists of prophase II, metaphase II, anaphase II and telophase II. Meiosis... 2
  • 5. PHASES OF MEIOSIS • Before entering meiosis I, a cell first go through interphase. • The interphase preceding meiosis is important because replication of DNA takes place during this stage. • Replication is confined to the synthesis (S) phase of interphase. • It is preceded by a post-mitotic gap phase (G1) and followed by a pre-mitotic gap phase (G2). • During these two phases there is no replication of DNA. PRE-MEIOTIC INTERPHASE 3 Meiosis...
  • 6. FIRST MEIOTIC DIVISION (MEIOSIS-1) • Cell division that reduces the chromosome number by one-half. • It separate homologous chromosomes and produce two cells with haploid chromosome number (n). • For that reason it is known as Reductional Division. • Four phases: Prophase I, Metaphase I, Anaphase I,Telophase I PROPHASE -1 • Longest and most complex phase. • It consists of 5 substages : Leptotene, Zygotene, Pachytene, Diplotene, Diakinesis. • In G2 and prophase of meiosis I, the two replicated chromatids of each chromosome are associated with each other by cohesin complexes. 4 Meiosis...
  • 7. LEPTOTENE • Chromsomes become very thin • They are not distinct, cannot be easily seen. PROLEPTOTENE • During this stage the chromosomes become more distinct. • Chromosomes appear as slender threads bearing a series of granule- like structures called chromomeres. • Under electron microscope the leptotene chromosomes have an axial filament. • To this filament, chromatin fibres are attached as a series of lateral loops. • Cytoplasm has many polyribosomes, but few ER. • A pre-meiotic pairing may takes place during leptotene. 5 Meiosis...
  • 8. • Leptotene chromosomes may be irregularly arranged, or may be polarized towards the centrioles forming a ‘bouquet’. • In plant cells the chromosomes may sometimes form a tangle of threads, called the synizetic knot, on one side of the nucleus. • During leptotene the cytoplasm has many polyribosomes, but endoplasmic vesicles are few. • A pre-meiotic pairing may takes place during leptotene. • Homologous chromosomes may be aligned together, but no synaptonemal complexes are formed. 6 Meiosis...
  • 9. ZYGOTENE • During zygotene the chromosomes become shorter and thicker. • The homologous chromosome starts pairing together. • These homologous chromosome come to lie side by side in pairs & this pairing is called synapsis. • The complex formed by a pair of synapsed chromosome is called bivalent. • Pairing may begin at the centromere and proceed towards the ends (procentric pairing). • They may begin at the ends and proceed towards the centromere (proterminal pairing). 7 Meiosis...
  • 10. • Sometimes it may occur simultaneously at all chromomeres (intermediate condition). • In many plant and animal species a kind of pseudosynapsis between non- homologous regions also occurs. • When synapsing occurs, the chromosomes do not lie side by side but rather on top of each other with a protein called the synaptonemal complex holding them together. 8 Meiosis...
  • 11. SYNAPTONEMAL COMPLEX • The synaptonemal complex is a proteinaceous structure that is formed between paired homologous chromosomes. • It is a tripartite ribbon like structure. • It consists of a dense central element, with a dense lateral element on either side. • The lateral elements are attached at both ends to the nuclear membrane. • The lateral elements are connected to the central element by a series of lateral loops which form transverse units or LC fibres. 9 Meiosis...
  • 12. • Three specific components of the synaptonemal complex have been characterized: SC protein-1 (SYCP1), SC protein-2 (SYCP2), and SC protein-3 (SYCP3). • Lateral elements are mainly formed by SYCP3 and secondarily by SYCP2. • Central element and transverse filaments are composed by the protein SYCP1. • The axial filaments of leptotene chromosomes later become transformed into the lateral elements of the SCs. • The SCs first appear during zygotene. • They are completed by the beginning of pachytene. • By diplotene the SCs have mostly disappeared, but remnants may be seen even in diakinesis. • They are believed to be involved in chiasmata formation and crossing over. • It stabilise the two homologous chromosomes till the crossing over is completed. Meiosis... 10
  • 13. PACHYTENE • The bivalents appear as thick threads due to condensation of chromosomes. • Two chromatids belonging to different homologues undergo one or more transverse breaks at the same level. • The breaks never occur between two sister chromatids. • The break is followed by interchange and fusion of broken ends between two homologous chromosomes. • This is called crossing over. • The site where crossing over occurs forms a recombination nodule. • An enzyme called recombinase is involved during this phase. 11 Meiosis...
  • 14. DIPLOTENE • Separation of paired homologues chromosomes begin. • The separating chromosomes are held together at one or more points called chiasmata. • A chiasma formed at the ends of chromosomes is called a terminal chiasma. • Chiasmata formed along the lengths of chromosomes are called interstitial chiasmata. • Terminalization of chiasmata takes place. • The synaptonemal complexes mostly disappear during diplotene. • Cytoplasm shows an increase in the number of endoplasmic vesicles. Meiosis... 12
  • 15. DIAKINESIS • The chromosomes become more contracted. • The bivalents are more evenly distributed in the nucleus and migrate towards the periphery. • The homologues remain in contact with each other by their terminal chiasmata. • The nucleolus is detached from the chromosome, or disappears. • Remnants of the synaptonemal complexes may still be seen. 13 Meiosis...
  • 16. METAPHASE-1 • The chromosomes now become arranged on the equator of the cell. • The spindle is formed. • Spindle fibres become attached to the centromeres of the two homologous chromosomes. • The two centromeres of each bivalent lie on opposite sides of the equatorial plate. • The nuclear membrane disappears. • The chromosomes reach their maximum contraction. • Spindle formation begins. PROMETAPHASE 1 Meiosis... 14
  • 17. ANAPHASE-1 • Separation of the two homologous chromosomes of each bivalent begins. • One chromosome from each bivalent begins to migrate to one pole, the other migrates to the opposite pole. • The sister chromatids are said to be co-oriented. • The kinetochores of each bivalent attach to spindle microtubules from opposite spindle poles. • They are said to be bi-oriented. • During Anaphase I original chromosomes separate, so reduction in the number of chromosomes from 2n to n number. 15 Meiosis...
  • 18. TELOPHASE-1 • The homologous chromosome complete their migration to the two poles because of shortening of spindles. • When the chromosomes reach the poles they persist for some time in the condensed state. • Later they undergo despiralization and become elongated. • The nuclear membrane is reformed, but the nucleoli do not reappear. • Unlike in mitosis the spindle fibres do not disappear completely. 16 Meiosis...
  • 19. CYTOKINESIS • Cytokinesis involves the formation of a cleavage furrow, resulting in the pocketing of the cell into two cells. • In most plant cells daughter cells are formed by the laying down of cell plate between the two groups of chromosomes. • In other plant cells cytokinesis does not take place until both meiotic divisions are completed. • At the end of Telophase I and Cytokinesis, two daughter cells are produced. • Each daughter cell with one half of the number of chromosomes of the original parent cell. Meiosis... 17
  • 20. 18 Meiosis... MEIOSIS I STAGES MAIN EVENTS Prophase 1 : Sub-Stages :- Leptotene Chromosomes appear as slender threads & Bouquet Formation Zygotene Synapsis & synaptonemal complex formation Pachytene Chromosomes appear as thick threads & Crossing over Diplotene Chiasma formation Diakinesis Chromosomes get thick and contracted Metaphase 1 Chromosomes arranged in equator region & Spindle formation Anaphase 1 Separation of homologous chromosomes & Beginning of chromosomes migration to opposite poles Telophase 1 Migration of homologous chromosomes complete & Shortening of spindle fibres
  • 21. INTERKINESIS OR INTRAMEIOTIC INTERPHASE • Interkinesis is a metabolic phase between telophase-1 & prophase-2. • During this phase chromosomes are elongated but do not form chromatin fibres. • This differs from interphase I and interphase of mitosis in that there is no duplication of DNA. • The RNA and protein required during meiosis-2 are syntesised during this phase. 19 Meiosis...
  • 22. MEIOSIS -2 • During Meiosis II, two sister chromotids of each chromosome separate and migrate to the opposite pole. • As a result, the number of chromosomes in each of the two haploid nuclei remains the same (i.e haploid), at the end of this division. • Thus the second division of meiosis is often referred to as equational division. • Sometimes, it is called as ‘Meiotic Mitosis’. • The second meiotic division is also divided into four stages : Prophase II, Metaphase II, Anaphase II, Telophase II. • The 4 stages of meiosis – II is same as mitosis. • It differs from mitosis in that DNA does not duplicate, while the centromeres do so. Meiosis... 20
  • 23. 21 Meiosis... MEIOSIS II STAGES MAIN EVENTS Prophase 2 Chromatids with widely separated arms & Spindle formation and nuclear membrane disappearance Metaphase 2 Chromosomes oriented at equatorial plate & Spindle fibre attachment as same as in mitosis Anaphase 2 Centromeres divide & Two chromatids of each chromosome separate to opposite poles Telophase 2 Reconstruction of nuclei. Nucleus, centriole and chromosomes return to interphasic condition & Nucleus contains haploid no. of chromosomes Cytokinesis Dyad divides into two parts. One parent cell produce four haploid daughter cells, together known as tetrad
  • 24. REMOVAL OF COHESION • To facilitate two consecutive chromosome segregation phases, cohesins have to be lost from chromosome. • During meiosis, cohesins are lost from chromosome arms by the end of meiosis I, but a pool of cohesins around kinetochores is protected from removal. • This pool of cohesins persists throughout meiosis I but is removed at the onset of anaphase II. • The mechanisms that remove cohesins during meiosis are the same as during mitosis. • Securin degradation releases separase, which then cleaves the cohesins. • This allows the recombined maternal and paternal chromosomes to separate, but each pair of chromatids remains associated at the centromere. • During metaphase II, sister chromatids align on the metaphase II spindle and separase is activated yet again, cleaving the residual cohesin around centromeres. 22 Meiosis...
  • 25. Rec8 • A specialized cohesin subunit, Rec8, is necessary for the stepwise loss of cohesins from chromosomes during meiosis. • Rec8 is homologous to Scc1, the cohesin subunit that closes the cohesin ring in the cohesin complex of mitotic cells. • During early anaphase of meiosis I, Rec8 is lost from chromosome arms but is retained at centromeres. • During early anaphase of meiosis II, centromeric Rec8 is cleaved by separase, so the sister chromatids can segregate. • The mechanism that protects Rec8 from cleavage at centromeres during meiosis I is similar to the mechanism that protects Scc1 at centromeres during mitosis. Meiosis... 23
  • 26. PP2A & SHUGOSHIN • Cohesion at the centromeres is maintained because a specific isoform of protein phosphatase 2A (PP2A). • PP2A is localized to centromeric chromatin by members of a family of proteins known as the Mei-S332/Shugoshin. • PP2A keeps cohesin in a hypophosphorylated state that does not dissociate. • During metaphase II, Mei-S332/Sgo1 dissociates from chromosomes. • When the last kinetochore is properly associated with spindle microtubules, Cdc20 is depressed and associates with the APC/C. • This causes ubiquitinylation of securin. • This releases separase activity, which cleaves Rec8, eliminating cohesion at the centromere and allowing chromatid separation in anaphase. 24 Meiosis...
  • 27. • Cohesin removal differs for meiosis I because when Rec8 replaces Scc1 in the cohesin complex. • The meiotic cohesin complex can only be removed from chromatin via the action of separase. • Rec8 also differs from Scc1 in that it must be phosphorylated by several protein kinases to be cleaved by separase. • During meiosis I, the centromere-specific isoform of PP2A targeted to centromeric chromatin by Mei-5332/Shugoshin prevents this phosphorylation. • The PP2A targeting factor and PP2A then dissociate from chromosomes by metaphase II, allowing separase cleavage of Rec8. Meiosis... 25
  • 28. • In mitosis and meiosis ll, sister kinetochores attach to spindle microtubules emerging from opposite spindle poles. • These kinetochores are said to be bi-oriented. • This is essential for segregation of sister chromatids to different daughter cells. • In contrast, at meiosis I metaphase, sister kinetochores attach to spindle microtubules emerging from the same spindle pole. • These sister kinetochores are said to be co-oriented. • Attachment of sister kinetochores to the proper microtubules in meiosis I and II is critical for correct meiotic segregation of chromosomes. ORIENTATION OF KINETOCHORES 26 Meiosis...
  • 29. • Monopolin complex is a protein complex that links with sister kinetochores during meiosis I for their attachment to microtubules emerging from the same spindle pole. • In organisms where kinetochores attach to multiple microtubules, Rec8- containing cohesins are essential for sister kinetochore co-orientation. • These meiosis-specific cohesins impose a rigid kinetochore structure. • This restricts the movement of sister kinetochores and thereby favoring attachment to microtubules from the same spindle pole. • Like during mitosis and meiosis II, correct attachment of meiosis I chromosomes is mediated by a tension-based mechanism. • During meiotic metaphase I, kinetochore-associated microtubules are also under tension. Meiosis... 27
  • 30. • Because chiasmata generated by recombination between homologous chromosomes and the cohesins distal to the chiasmata prevent them from being pulled to the poles. • Kinetochore-microtubule attachments are unstable in the absence of tension. • When kinetochores attach to the wrong spindle fibers release the incorrect microtubules. • This enables them to bind microtubules again until attachments are made that generate tension. • As in mitosis, once tension is generated, microtubule attachment to the kinetochores is stabilized. 28 Meiosis...
  • 32. REFERENCE • Burgess, J. (1985). Introduction to plant cell development. CUP Archive. • Lodish, H. (2008). Molecular Cell Biology. New York: W.H. Freeman and Company. • Pawar, C.B. (1983). Cell Biology. Himalaya Publishing House. • https://www.slideshare.net/KAUSHIKKUMARPANIGRAH/presentation-on-meiosis • https://www.google.com/amp/s/slideplayer.com/amp/14961858/ • https://www.khanacademy.org/science/ap-biology/heredity/meiosis-and- genetic-diversity/a/phases-of-meiosis 30 Meiosis...