Linkage and

crossing over

Dr. TARUNPREET SINGH

LINKAGE CROSSING OVER
• Discovery of linkage • Discovery of crossing over
• Meaning of linkage • Meaning of crossing over
• Characteristics of linkage • Characteristics of crossing
• Genes in linkage over
• Theories • Types crossing over
• Kinds of linkage • Mechanisms
• Linkage group • Factors affecting crossing
• Significance over
• Significance
• Differences between
crossing over and linkage
 Discovery of Linkage
• The principle of linkage was
discovered by English Scientists
William Bateson and R.C.
Punnet in 1906 in Sweet Pea
(Lathyrus odoratus). However, it
was put forward as a regular
concept by Morgan in 1910
from his work on (Drosophila
melanogaster).
 Meaning of Linkage
• Linkage is the phenomenon of AB

certain genes staying together

during inheritance through
AB
several generations without any
change or separation due to their
being present on same
chromosomes.
 CHARACTERISTICS OF
LINKAGE
• Linkage involves two or more genes which are linked
in same chromosomes in a linear fashion.
• Linkage reduces variability.
• It may involve either dominant or recessive
alleles(coupling phase) or some dominant and some
recessive alleles(repulsion phase).
• It usually involves those genes which are located close
to each other.
• The strength of linkage depends on the distance
between the linked gene.
*Lesser the distance higher the strength of linkage*
 Genes in Linkage
• LINKED GENE :
These genes do not show
independent assortment.
It occurs in same
chromosome.
Dihybrid ratio of linked
gene is 3:1
• UNLINKED GENE:
These gene show
independent assortment.
Dihybrid ratio is 9:3:3:1.
 Theories of Linkage

• DIFFERENTIAL MULTIPLICATION
THEORY (William Bateson)

• CHROMOSOMAL THEORY (Thomas Hunt

Morgan)
 DIFFERENTIAL
MULTIPLICATION THEORY

• This theory was put forward by Bateson in

1930.
• He said that after the segregation of genes
during gametogenesis certain genes multiply
faster then others.
 CHROMOSOME THEORY
Morgan and Castle associate genes with
chromosome and formulated postulates
The genes which show Linkage are located in
same chromosome.
The distance between linked gene in the
chromosome determine the strength of linkage.
The genes lie in a linear manner in the
chromosomes.
 KINDS OF LINKAGE
ON THE BASIS OF CROSSING OVER
i. Complete linkage
ii. Incomplete linkage
ON THE BASIS OF CHROMOSOME
INVOLVED
i. Autosomal linkage
ii. Allosomal /Sex linkage
ON THE BASIS OF GENE INVOLVED
i. Coupling phase
ii. Repulsion phase
 ON THE BASIS OF CROSSING
OVER
 COMPLETE
LINKAGE
• The genes located in
the same chromosome
are inherited together
over the generations
due to absence of
crossing over. It is rare
but has been reported
in male Drosophila.
 INCOMPLETE LINKAGE
• Genes present on
the same
chromosomes have
a tendency to
separate due to
crossing over. They
produce
recombinant
progeny beside the
parental types.
 ON THE BASIS OF
CHROMOSOMES INVOLVED
 Based on the chromosomes involved:- Based
on the location of the genes on the chromosomes,
linkage is categorized into:-
i. Autosomal linkage:- It refers to linkage of
those genes which are located in autosome
(other than the sex chromosomes).
ii. Allosomal linkage:- It refers to linkage of genes
which are located in sex chromosomes i.e. either
“X’or “Y’.
 ON THE BASIS OF GENES
INVOLVED
• Depending on whether all dominant or some dominant and recessive alleles are
linked together, linkage can be categorized into coupling and repulsion phase:-
i. Coupling phase:- Dominant alleles and recessive alleles present on the same
chromosomes shows coupling phase.
TR tr
----- ---- COUPLING PHASE
TR tr
ii. Repulsion phase:- Dominant alleles of same genes are linked with
recessive alleles of other genes on same chromosomes shows repulsion phase.
Tr tR
---- ---- REPULSION PHASE
Tr tR
 Linkage group
• A linkage group is a linearly
arranged group of linked
genes which are normally Linkage group
inherited together.
• Example:- In a fruit fly
Drosophila melanogaster has
four linkage group (Four
pairs of chromosomes).
• In human being 23 linkage
group are present (23 pairs of
chromosomes).
 Significance of linkage
• It reduce the chances of formation of new
combinations of genes in gametes.
• It helps keeping the parental, racial and
specific traits together.
• It also useful for maintaining the good
character of newly developed variety.
• Linkage plays an important role in determining
the nature and scope of hybridization.
CROSSING OVER
 Discovery of crossing over
Frans Alfons Janssens who
described the phenomenon
of crossing over in 1909.
He is observed cross-like
arrangements in meiosis
and proposed crossing over
as a genetic process.
 Meaning of Crossing over
• Crossing over or (chromosomal cross over) is
the exchange of genetic material between
homologous chromosomes that results in
recombinant chromosomes.
 Characteristics of crossing over
 Crossing over occurs between non-sister chromatids. One
chromatid from each of the two homologues chromosomes
is involved in crossing over.
 Crossing over leads to re-combinations or new combinations
between linked genes.
 The value of crossover or recombinants may vary from 0-
50%.
 Crossing over generally yields two recombinant types or
crossover types and two parental types or non-crossover
types.
 Crossing over generally leads to exchange of equal segments
or genes and recombination is always reciprocal.
 Stage at which crossing over occur
• The meiotic crossing over
takes place during the
pachytene stage of the
prophase of meiosis –I.
Pachytene stage is also
known as recombination
stage. Crossing over
occurs when homologus
chromosomes are in the
four chromatid or tetrad
stage in pachytene.
 Types of crossing over
It is mainly two types
 Somatic or mitotic crossing over.
 Germinal or meiotic crossing over. It is further
divided into two types
• Equal crossing over
• Unequal crossing over
 Somatic or mitotic crossing over

• This type of crossing over occurs in the

somatic cells during mitosis.
• It is rare and has no genetic significance.
• Example- Curt Stern reported it in the fruit
fly and Potnecorvo noted it in the fungus
Aspergillus
 Germinal or meiotic crossing over
• This type of crossing over takes place in the
germinal cells during meiosis that produces
gametes.
• It is universal and has a great genetic
significance.
Kinds of Germinal crossing over
• (A) Equal crossing over :- The segments
exchanged between the chromosomes are of
equal size. It is divided into three types
according to the number of points at which it
occurs.
 Single crossing over
 Double crossing over
 Multiple crossing over
a) Single crossing over In
this type of crossing over
the chromatids break and
reunite at one point only.
b) Double crossing over
During this type of
crossing over the
chromatids break and
reunite at two points in the
same tetrad.
c) Multiple crossing over: In
multiple crossing over,
chromatid break and
reunite at many points in
the tetrad . It occurs rarely.
Unequal crossing over
• The segments exchange between chromatids
are unequal so that one chromosome
receives an extra gene, and other gets one
gene less.
 MECHANISM OF CROSSING
OVER
I. Synapsis
II. Tetrad formation
III. Exchange of chromatid segments
IV.Terminalization
 1.Synapsis
• In the Zygotene
or pairing stage of
prophase-I, the
homologous
chromosome of
each pair come
together and line
up side by side.
This pairing of
homologous
chromosomes is
called synapsis.
 2.Tetrad formation
• The two chromatids of
chromosome are referred to as
dyad. A group of four homologous
chromatids (two dyad) of two
synapsed homologous chromosome
is known as tetrad. The two
chromatids of same chromosome
are called sister chromatids. The
two chromatids, one of the one
chromosome and other of its
homologue, are termed non-sister
chromatids.
 Synaptonemal complex
• A highly organized structure of
filaments is formed between the
paired homologous chromosome at
the zygotene and pachytene stages
of meiosis-I, the structure is called
synaptonemal complex. It helps in
crossing over by keeping the
homologous chromosome in
closely paired state.
 3. Exchange of chromatid segments
• Two non sister chromatid in a tetrad break at
equivalent locations.
• The broken ends transpose and join the
respective broken ends of opposite chromatid.
• This complete the process of crossing over.
• The unchanged chromatids are called parental
or non cross overs.
• The changed chromatids are called
recombinants.
3. Exchange of chromatid segments

Physical breakage
of chromatids into
segments

Transposition
of segments

Reunion of segments
 4.Terminalization
• Completion of crossing over marks the end of
pachytene stage and beginning of diplotene stage.
• Synaptic forces end and the homologous
chromosomes separate.
• The points at which the separation does not occur
is called chiasmata.
• The chromatids separate progressively from the
centromere towards the chiasma which moves
like a zipper towards the end of tetrad.
• The slipping of chiasmata towards the ends of the
bivalents is called terminalization.
Terminalization
 Factors affecting crossing over
Maternal age effect
Temperature
Nutritional and chemical effect
Chromosomes effect
Centromere effect
Mutation effect
Sex
 Significance of crossing over
• Crossing over has helps in establishing the
concept of liner arrangements of genes.
• The frequency of Crossing over helps in
mapping of chromosomes i.e. determining the
location of genes on the chromosomes.
• It is an important factor in sexual reproduction.
• It increases the variation which is vital for
evolution.
• It helps in plant breeding also.
 Difference between Linkage and
Crossing over
Linkage Crossing over
1. It keeps the genes together. 1. It leads to separation of linked
genes.
2. It involves exchange of
2. It involves individual segments between non-sister
chromosomes. chromatids of homologous
chromosome.

3. The number of linkage 3. The frequency of crossing

groups can never be more over can never exceed 50%.
than haploid chromosome
number. 4. It increase variability by
4. It reduces variability forming new gene
combinations.
 REFERENCES
Kohli J (2002) Linkage and crossing over Inst.
Cell Biol Univ of Berne, Berne, Switzerland.
K.N Bhatia and Neelam Dhand (2014) Cell
Biology and Genetics Trueman Publication, New
Delhi.
P.S Dhami and J.K Dhami (2015) Text book of
Zoology Pradeep’s Publication Jalandhar (India).
M.P Arora and G.S Sandhu (2000) Genetics
Himalaya publication Bombay.