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Meiosis has a number of overall functions
dÊ ’educe the chromosome number by half from diploid(2n) to haploid(n).
dÊ ach gamete nuclei will contain one chromosome from every homologous pair.
dÊ 3ncrease genetic diversity.
This section assumes that the information in section 4.2. Student should review that work
before adding the following details.
Additional notes are provided for HL students.
dÊ Meiosis consists of two rounds of cell division.
dÊ M1 : ’eduction division which separates the chromosomes of a homologous pairs.
ach gametic cell will contain one of the two chromosomes from every homologous
pair. This has great genetic significance since it separates the alleles for every gene.
dÊ M2: Separation of the 'sister chromatids'. After M1 the gametic cells contain a pairs
of 'sister chromatids'. The separation of the 'sister chromatids' also produces
variation in gametes since this will isolate the 'recombinants alleles' due to cross-
over in a gamete cell.

Meiosis is proceeded by the interphase which includes the replication of DNA (S-Phase) so
that when meiosis begins, it starts with each chromosome represented by a pair of 'sister
chromatids' held together by a centromere.
@c    

› ^
›
:
dÊ hromatin begins to condense with super coiling reaching it peak(x16, 000) in
metaphase 3.
dÊ A process called synapsis which brings together the chromosomes of a homologous
pair (do not confuse with sister chromatids) begins. This only lasts until cross-over
has occurred. After cross-over has completed in metaphase 3 the homologous pairs
are held together only at the chiasmata.
dÊ The chiasmata are the positions of DNA exchange which is called cross-over.
dÊ The non-sister chromatids of a homologous pair adhere along the length of the
chromosomes.
dÊ The four chromatids from each pair of homologous chromosomes are called a tetrad
or bivalents.
dÊ ross-over occurs during the prophase but is not visible until the final condensation
at metaphase.
dÊ During the condensation of the chromosomes, a point is reached when the
homologous pair seems to repel each other except at regions called chiasma. The
centromere's of one pair of 'sister chromatids' particularly repels the centromere
region of the other in the homologous pair.
dÊ rossing over increases the genetic diversity of the gametes and therefore increases
variation in successful fertilizations. The process of forming µrecombinants¶ is dealt
with in section 10.2 µgene linkage¶.
dÊ Nuclear membrane breaks down and the nucleoli disintegrates.
dÊ 3n animal cells the centrioles are placed at either pole of the cell and serves as a
focal point for the organisation of the spindle microtubules.
dÊ Microtubules attach to the centromere's of each pair of homologous chromosomes.


 À ››

dÊ Homologous pairs are aligned on the
equatorial plate.
dÊ Homologous pairs have been held together
by the chiasmata but this is not shown for clarity.
dÊ ach 'pair of sister chromatids' in a
homologous pair is attached to microtubule. ach
half of the spindle is attached to the opposite poles.

À
 ››

dÊ The spindle microtubules contract pulling the
pair of 'sister chromatids' to one pole and the other pair of 'sister chromatids' for the
same homologous pair to the other pole.
dÊ 3t seems that this process is a combination of spindle contraction (75%) and a
centromere motor (25%) pulling the pairs of chromatids along the spindle.
dÊ At this point the chiasmata break down and the exchange of lengths of DNA including
alleles of genes is complete.

À 

›

dÊ The genetic material is now organised at the two pole of the cell.
dÊ ach pole contains a pair of 'sister chromatids' , one from each homologous pair.
dÊ 3n some species the nuclear envelop re-forms whilst in others there is an immediate
progression to M2 and prophase 33.
@         

À ^
›

dÊ hromosomes condense again if required.

There has been no chromosome replication but each chromosome is represented by

a pair of 'sister chromatids'.
dÊ The spindle forms in a new plane for each of the two cells. The positioning of the
plane for the spindle is critical for later development.
dÊ The 'sister chromatids' attach to the spindle microtubule at the region of their
centromere.
dÊ The 'pairs' then move around, beginning to align for the metaphase 33.

À ››

dÊ The pairs of sister chromatids line up at the
equator of each cell.
dÊ The are now very condensed and at their most
visible.
dÊ The cohesin joining together the arms of the
chromatids has broken down except at the centromere.
The chromosome takes on the characteristic open X -
shape.
dÊ ach of the chromatids will separate at the next
stage.

À  ››

dÊ The
combination of
spindle contraction and the centromere 'motor'
pulling on the spindle microtubule, pulls
the sister chromatids apart.
 dÊ ëne chromatid from each pair goes to each pole.
dÊ ach chromatids can now be referred to as a chromosomes again.
dÊ The chromosomes from the pair of 'sister chromatids' are not identical due to the
exchange of DNA during cross-over (not shown in diagram).
À

›

dÊ The chromosomes gather at opposite pole and form the new haploid nuclei.
dÊ The nuclear envelope material that has remained within the cytoplasm re-forms
around the sets of chromosomes.
dÊ There are now two nuclei per cell, each is haploid
À  
 


dÊ The cells divide to form a tetrad of four haploid nuclei.
dÊ The type of division depends on this being an animal or plant cell.
3n the sexually mature human male the process of meiosis and the production of mature
motile sperm cells (haploid) takes over a month. 3n the human female the process begins
during embryonic development but then undergoes a period of 'dormancy'. The cell still
diploid is held at prophase 3 until ovulation many years later. The completion of meiosis
does not actually end until fertilisation and then in humans this results in a single gametic
cell and not the four as described above. (see reproduction unit). ëther species have refined
this process in different ways as an adaptation to their own ecology.

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(a) 3nterphase:
During the interphase each of the chromosomes in a
homologous pair replicates. The two copies of a
chromosome are held together by a centromere. The
replicated chromosome pair are described as 'sister
chromatids'.

(b) Prophase 3:
Molecules, 'cohesin's', hold the homologous pairs close
together. This facilitates the homologous pair joining to
the same spindle microtubule. The exchange of DNA
between parallel arms of the non-sister chromatids
takes place.
(c) Prophase 3 -Metaphase 3:
Still in prophase the DNA molecule exchanges length of
DNA.
3n metaphase the chiasmata are more obvious as the
'cohesin's' are broken down. The homologous pairs seem
to repel each other particularly at the centromere.
However they remain linked together at the chiasma.
This is also the peak phase of condensation.
(d) Anaphase 3:
The separation of the homologous pairs in the anaphase
finally breaks the chiasma connections.
The DNA exchange on the arms of the chromosomes is
complete
(e) Anaphase 33:
3n anaphase 33 the 'sister chromatids' are separated into different cells.
Some of the chromosomes are new recombinants of DNA containing part maternal and part
paternal chromosomes.
3n effect this has created new combinations of linked genes.
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§ 
› ›
 ›






 
dÊ 3ncreasing genetic diversity in gametes which in turn increases genetic diversity of
the populations.
dÊ rossover as described in the previous section creates new combinations of linked
genes.
dÊ This creates new genotypes for the gametes that are not due to random assortment.


















› 

 ›

At Metaphase 3, homologous pairs aligned on the equatorial plate of the dividing cell. The
diploid cells are the centre row. They will divide, as for anaphase 3,vertically.
The homologous pair are both held on the same spindle microtubule (green dashed line).
Anaphase 3 will separate the homologous pair and therefore their alleles.
However with this cell (diploid number = 4) then there are two possible orientations of the
homologous pairs on the equatorial plate.
’ANDëM orientation means that all orientations are equally possible.
3n this example the number of possible gametes is 4.
3n general the calculation of the number of possible gametes = 2n
Homo sapiens 2n=46 , n= 23 therefore number gametes = 2n = 8,388,608.

§ 
› ›

ombining cross-over with random orientation creates a great deal of variation. The greater
the diploid number and the greater the degree of cross-over, the greater the diversity.
There are other genetic factors that increase the genetic variation of a population still
further:
 dÊ ’andom fertilisation in which any possible gamete form the male (2n) can randomly
fertilise any other possible gamete from the female (2n) = 22n.
dÊ §ene mutation
dÊ hromosome mutation
Populations are subject to other means of increasing genetic variation. Students might like
to consider what these might be, think about the definition of a population and the factors
that affect population size.

›

› 



dÊ This topic reveals the source of genetic variation. That is the source of variation on
which natural selection acts. The random orientation, cross-over and mutation are
random processes. The natural selection of a particular phenotype is not.
dÊ This topic requires us to regard sexually reproducing populations as genetically
diverse and that diversity within the population changes from one generation to the
next.
dÊ Living things possess a genetic program which distinguishes them from other types
of matter.
dÊ The population of a species shows genetic varation. Knowing this makes it difficult to
be specific about how a population will respond to a change in their environment.
Therefore biology requires a 'population thinking' approach so that phenomena might
be understood. This is what distinguishes Biological Science from the Physical
Sciences.

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3n his work Mendel followed patterns of inheritance of characteristics in

plants. He is perhaps most famous for his work on pea plants from which he
developed and published his laws of inheritance.

Law of 3ndependent Assortment:

dÊ This law states that allele pairs separate independently during the formation of
gametes.
©Ê Therefore, traits are transmitted to offspring independently of one another.

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! @  !     
  

Mendel's law of independent assortment has been covered in the previous section 10.1.3 by
discussing the random orientation of the chromosomes at metaphase 3. We should
remember that Mendel tracked the tracked the inheritance of physical characteristics and
traits (phenotypes) not the actual alleles or chromosomes.
Mendel's law of independent assortment>
››

› ›      
›

›
.
3n terms of meiosis:
› 

 ›












 ››


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Terminology:
dÊ 0 




involve two genes which control two characteristics. There are
complications of these patterns as illustrated in the calculations that follow.
 dÊ r   genes are found on different chromosomes and can be segregated by
random assortment of meiosis/ metaphase 33
dÊ 


› are those chromosomes other than the XY gender determining
chromosomes (not sex linked).
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!   (

3n some species there is one homologous pair which carry genes that determine gender.
THis work has already been covered in the section onsex chromosomes.
There are other genetic conditions (also covered in topic 4) associated with the so called
'sex linked" conditions.

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!      
 "   (   

’ead the next section 10.2.4 first and understand the concept of a linkage group which is
key to understanding of this particular assessment statement.

The diagram shows the loci of genes A and B.

The alleles A is linked to B, a is linked to b.
The homologous pairs are held together like this
during prophase 3.

The crossover point occurs above the loci for

gene A and below the gene loci for A.
The position at which the exchange occurs is
called the chiasma.
The recombinants will form between non-sister
chromatids which are crossing over.
The homologous pairs remain attached at the
chiasma until anaphase 3 when they are pulled
apart.

After anaphase 33, the chromatids are separated.

The linked genes are:
dÊ A with B
dÊ a with b
dÊ recombinant a with B
dÊ recombinant A with b

§ 






dÊ The genotype of the gametes from
meiosis above would be AB and ab.
dÊ After cross over there would also be Ab and aB.
However most cells of the cell undergoing meiosis to form a gamete will not cross over
between these two points.
Therefore:
dÊ §ametes Ab and ab will be high frequencies.
dÊ §ametes Ab and aB will be low frequency.
When cross over occurs there will be low frequency of recombinants produced.
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The genotypes of an organism is AaBb. This looks like a dihybrid such as those in the
sections above. The two genes would be on separate homologous pairs, however there is an
alternative and this is known as linkage.
These two genes are actually on the same chromosome.
The  allele is linked to the  allele along the length of DNA.
The › allele is linked to the allele along the length of DNA.
3n the image take after S-phase, the homologous pair is aligned as it
would be found in Prophase 3 of meiosis.
3t might be predicted that during gamete formation of fertilisation that
the A allele will be followed by the B allele. This would be more like
carrying out two simultaneous monohybrid crosses.

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xample: Sweat Peas (Ô

 
) Ê
This famous example comes from the
work of W. Batson who rediscovered
the work of Mendel and carried out
his own experiments in the early 20
entury.
Allele Key:
dÊ ülower colour Purple (P) and ÊÊÊÊ
’ed (p)
Ê
dÊ Pollen grain shape, Long (L)
and short (l)
A cross was made between a plants
that where heterozygotes at both
gene loci (PpLl).
When dealing with linked genes the
linkage group is drawn on the
horizontal with a line representing the
chromosome. Using the above cross
the cross would be represented.

Parental phenotypes are the same PpLl

being heterozygous at both gene loci.

3f the gene for flower colour and pollen

grain shape are linked then the
genotypes are shown as left.

The gamete genotypes are shown to the left with the alleles for flower colour and pollen
grain shape shown linked by the horizontal line.

The possible recombinants from cross over are shown for one parent only.
These recombinant gametes will be at low frequency.




The grid shows the cross with the possible fertilisations in the boxes.
There four expected genotypes.
There are four recombinant genotypes but cannot be observed from phenotypes.
There are two recombinant phenotypes which would be actually observed during the
experiment.


›


 



The result of W Batson's cross showed
that the result are neither dihybrid and
unlinked (9:3:3:1)or linked with no
linkage (3:1)
The result shows a rough 3:1 ratio of
Purple/Long : ’ed/Short
There are also recombinant phenotypes
at much lower frequency.


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’ecombinants are recognised by:
dÊ Unpredicted combinations of characteristics. (’ed Long, Purple Short).
dÊ Low frequency of new combinations of phenotype.
dÊ Statistical significant difference from ratios expected from either dihybrid and
unlinked (9:3:3:1)or linked with no linkage (3:1)

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Definition: 'A single characteristic that is controlled by two or more genes'
Ê
ach allele of a polygenic character often contributes only a small amount to the over all
phenotype. This makes studying the individual alleles difficult.
3n addition environmental effects smooth out the genotypic variation to give continuous Ê
distribution curves.

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a) 3s the genotypic variation
in the population. The more
genes involved with the
characteristic the greater
the number of phenotypic
classes.
(b) Phenotypic variation =
genotypic variation + environmental variation. The environmental component smooth the
genotypic category differences.