Linkage and Crossing

over
Linkage
• According to Mendel’s Law of independent
assortment, two pairs of characters used in a
cross assort independently.

• But, Each chromosomes contains many genes

which will not assort independently rather they
are inherited together.
• The genes situated on the same chromosome are
called linked genes.

• The phenomenon of inheritance of linked genes

from one generation to another is called linkage.
In other words,

• The tendency of two or more genes of the same chromosomes to

remain together in the process of inheritance is called linkage.

• Linkages was clearly understood by T.H. Morgan (1910) on the

fruit fly (Drosophila melanogaster) as chromosomal theory of
linkage.

The chromosomal theory of linkage states:

a. Genes lie in the chromosome in the linear order.
b. The linked genes stay on the same chromosome.
c. The tendency of genes to remain together and to express
together like their parents is because of their presence on the
same chromosome.
d. The strength of linkage is directly related to the distance
between the linked genes on a chromosome. The closer the
genes, the stronger is the linkage.
• The genes present on different chromosomes are called
unlinked genes.
• Unlinked genes assort (inherited) independently.
• The distance between linked genes is measured in map units
or morganoid or centimorgan.
• 1 map unit= 1% recombination or centimorgan.
• Linkage is possible if the distance between linked genes is 50
mu or more.
• If the distance is less than 50 mu, there is no crossing over
and non-parental recombinants are not formed.
• If the distance is more than 50 mu, the formation of new
recombinants is possible due to crossing over.
Linkage group: Linked genes present in a group on the same
chromosome are called linkage group.
• Linkage group in a cell is equal to the pair of chromosomes
present in a cell of an organism.
• For example maize has ten linkage groups (10 pairs of
chromosome), fruitf ly has 4 linkage group, pea has 7 linkage
group and human has 23 linkage group.
• But in male human being, the linkage group is 24 (22+X+Y).
• The size of linkage group depends upon the size of the
chromosome.
• Linkage group of any species =N
• Linkage group of any male species=N+1

Linkage map: The linkage map is the map of genes present on a

chromosome based o n linkage analysis.
• A linkage map doesnot show the physical distances between
genes but their relative positions.
Significances of Linkage:
 The linked characters are preserved for many generations.
 It brings crossing over responsible for genetic variations in
organisms.
 It also reduces the possibility of variability in gametes
 It is useful for maintaining the good characters in new
variety.
 Inheritance of two linked genes controlling two different
characters will help in improvement of characters in the
offspring.

 Linkages was studied by T.H. Morgan (1910)by his

experiments in the inheritance of eye color in fruit fly
(Drosophila melanogaster). Red eye color is dominant
while the white eye colour is recessive.
Reasons for selecting Drosophila for Genetic study
 It can be cultured easily under a normal condition.
 It is harmless and inexpensive to culture.
 A large number of Drosophila can be kept in small space.
 Progeny produced after each mating is large.
 It breeds throught the year.
 Its life cycle is short and completes in 10-12 days, so the
result of controlled breeding are quickly available.

 The Drosophila has 4 pairs of chromosomes. 3 pairs are

autosome sand 1 pair is sex chromosome
 Male has 2 X chromosomes and female has One X and
One Y chromosome respectively.
Types of linkage: Linkage can be of
two types

1. Complete linkage

2. Incomplete linkage
1. Complete linkage

Complete linkage occurs when the genes for a

particular character are very close to each other that
they have no chance of separation from each other.

These genes always transmitted together from

generation to generation.

This phenomenon is called complete linkage.

• It is found in male Drosophila. Example is given below:
Red eyed normal winged Purple eyed vestigial winged
RRWW rrww

RrWw (hybrid/Red eyed and normal winged)

Test cross with homozygous recessive parents.

RrWw X rrww

RrWw RrWw rrww rrww

Red eyed normal wing (50%) Purple eyed vestigial wing (50%)
Phenotypic ratio- 1 : 1

The results indicate that red eyed character is linked with normal wings
while the vestigial wing is linked with purple eye.
The new recombinant does not occur as the parental genes are
completely linked with each other. The phenomenon is called
complete linkage.
2. Incomplete Linkage

The genes which are located far away from each other
show a tendency to separate during crossing over are
called incompletely linked genes.

The phenomenon of inheritance of these genes is

called Incomplete linkages.

• The incomplete linkage produce some percentage

of non-parental combinations (recombinants)
OR Example of incomplete linkage
Coloured and filled seeds Colour less and shrunken
CCSS ccss

CcSs (Colored and filled seed)

Test cross;
CcSs X ccss

CcSs Ccss ccSs ccss

Phenotypic ratio: 7:1:1:7

From the result it can be concluded that the linkage is incomplete and crossing
over has occurred to produce new recombinants than the parental
combination.

The phenomenon of this incomplete linked genes to produce mew

recombinant in f2 generation in test cross is called incomplete linkage.
 Sex linked inheritance

• Sex-Linked Inheritance is the inheritance of a trait (phenotype) that is

determined by a gene located on one of the sex chromosomes.
• The sex chromosome of animals and human beside having genes for sexual
characters also posses the genes for some non-sexual characters (somatic
characters) and they are carried together from one generation to another.
• Such genes for non sexual characters being linked with sexual genes in a sex
chromosome are called sex-linked genes and inheritance of such characters
are called sex-linked inheritance.
Sex linked inheritance
In human beings, there are two types of chromosomes,
autosomes and sex
chromosomes.

• Out of total 23 pairs of

chromosome 22 pairs are
autosomes and one pair is
sex chromosome.
• In females, the two sex
chromosomes are similar
X and X. But in male the sex
chromosomes X and Y are
morphologically different.
• The Y chromosome is J-shaped and is much smaller than the X
chromosome.
• Sex chromosome in male individual is heteromorphic
(different).
• The genes for sex linked characters occurs in either X-
segment or Y- segment or on both segments of
chromosome.
• The genes are either called X-Linked genes or Y-linked
genes for a particular trait. Based on location, sex linked
inheritances are of following types:
X-linked traits: Genes for these characters are located on
the non-homologous segments of X-chromosome. It
folllws criss-cross inheritance. Eg. Eye color in Drosophila
Y-linked traits: Genes for these characters are located on
the non-homologous segment of Y- chromosome.
X-Y linked traits: The genes for some characters located on
X and Y chromosomes are called X-Y linked genes. e.g color
blindness, skin cancer etc.
Characters of sex-linked traits (genes)
• Majority of sex linked traits are recessive.
• Male suffers from sex-linked disorders more often than
females.
• Mother passes the allele of sex linked trait (XX) to both
sons and daughters.
• Father passes the X chromosome to daughter always and
Y chromosome to the son.
• Only homozygous females can express a recessive trait,
while heterozygous females are carriers.
• In all sex linked traits heterozygous females are the
carriers.
• Since X-chromosome of son is always from the mother,
there is a 50% chance that the son would have the
disease trait.
• In other word, if a son gets a sex-linked disease, it is
always inherited from mother. (when both parents are
normal)
Criss-cross inheritance
It is a type of sex-linked inheritance where a parent passes the
traits to the grand child of the same sex through offspring of
the opposite sex.
i.e.
pass trait through son
Mother (carrier) -----------------------------------Grand daughter

Pass trait through daughter

Father (sufferer) -----------------------------------Grand son
• In criss-cross inheritance father does not pass the sex-
linked allele of a trait to his son. But the same is passed to
the daughter because the males have only one X-
chromosome which is transferred to the female offspring .
Only Y-chromosome of father is transferred to the male
offsprings but this sex chromosome does not carry many
alleles.
1. Criss- Cross Inheritance in Drosophila
While working on Drosophila, he noted sudden
appearance of one white –eyed male in the culture of red-
eyed Drosophila.
They have 4 pair of chromosome.
(sex chromosome:
XX in male and XY in female)
Carrier- Those individuals which carry the allele of trait
which might be passed on next generation.
They do not show symptoms, only carry the genes.

Sufferer- Those who carry the allele of trait responsible for

certain diseases which is expressed in them.

Type of eye color in Drosophila

Normal: red color eye
Mutant: white color eye
Example (Cross 1)
When Red Eyed female (XR XR)and White Eyed male (XwY) is crossed
all flies in F1 generation were found to be red.
Red Eyed female (XR XR) White Eyed male (XwY)
Gametes..XR XR Xw Y

XR Xw XR Y
F1 generation…Hybrid female (red) Red eyed male

When selfing the F1 generation;

On selfing
XR Xw XR Y

XR XR XR Y XR Xw XwY ………. F2
Red eyed female Red male red eyed female(hybrid) White eyed male
(hybrid)

All f2 females were red

Half of F2 males were white and half red eyed.
Red and white: 3:1 (phenotypic ratio)
A cross between Red eyed female and white eyed male:
• Morgan crossed a normal red eyed female with a white eyed male.
He found only red eyed males and female in F1 generation. When
the red eyed male and red eyed female individuals of F1 generations
are intercrossed, he got :
a.All F2 females were red eyed.
b.Half of F2 males were red eyed.
c.Half of F2 males were white eyed.
Thus, F2 generation shows red eyed and white eyed individuals in 3:1
ratio. The ratio of male and female flies was 2:2 or 50% each. Thus,
appearance of white eyed character is linked with the male sex, but is
inherited through the female. This shows criss-cross inheritance.
From the first cross he recognized that the eye color allele
were present only on X-chromosome and no homologous
allele present on Y- chromosome.

To verify this he further crossed F1 red eyed female hybrid

of F1 generation (XR Xw) and white eyed male (XwY)
Cross 2
F1 red eyed female (XR Xw) X white eyed male (XwY)

XR XW Xw Y

XR Xw XRY XwXW XW Y
Red eyed female Red eye male white eyed female white eyed male
It indicates white color is not restricted to male sex.
Cross 3 A cross between white eyed female and red-eyed male.
White eyed female & Red eyed male
Xw Xw XR Y

F1…………….XRXW XWY XRXW XWY

Red female (hybrid) White male red female white male

XR XW and XW Y are formed in F1 generation.

All females are red.
All males are white.

Now selfing of f1 generation,

 Red eyed female & White eyed male
XR Xw XW Y

F2…………….XRXW XRY XWXW XWY

Red female (hybrid) red eyed male White eyed female white eyed male
Cross between white eyed female and red eyed male:
The white eyed females were crossed with red eyed males. The result
that all males were white eyed, all females were red eyed in F1
generation. He got;
a.Half of F2 population were red eyed.
b.Half of F2 population were white eyed.
c.Half of F2 males were red eyed.
d.Half of F2 males were white eyed.

Morgan concluded that the eye color in Drosophila is a sex- linked

character and follows a criss-cross inheritance. The male transmits
sex-linked characters through his daughters to grandsons and never
through his sons.
2. Sex linked inheritance in human
Colorblindness in human
Color blindness is a defect in which a person cannot
distinguish red, green or both these colors from other
colors.
• It is recessive and X-linked trait. So it is only
transmitted through X-chromosome.
• Mutation causes defect in the red green pigment and
causes color blindness which then is inherited to
offspring following criss-cross inheritance.
• Colorblindness is a recessive and x-linked trait.
• Genes for these characters is located on non-homologous segment of
X-chromosome.
• Allele of this gene does not occur in Y-chromosome. So the trait is
transferred from father to his daughters and carrier mother to her son.

Symbols used in cross for color blindness

Colorblind female: XC XC
Carrier female: XC X
Normal female: X X
Colorblind male XC Y
Normal male: X Y
Cross 1
When color blind man marry a normal visioned woman

XCY x XX……………………Parents

XC Y X X…….Gametes

XCX XCX XY XY……..F1

Daughters are carrier while sons are normal.

When F1 daughter married to normal man, 50% of sons are color blind
while 50% are normal and all daughters are normal, while 50% of them
are carrier.

F1 carrier female Normal male

XCX x XY

XC X X Y

XCX XCY XX XY……………F2

Carrier daughter Color blind son normal daughter normal son

The character of carrier female received from her father is transferred to

her son.
When a normal visioned woman marries a color blind man,
all their male and female children will have normal vision,
but the female children are heterozygous, who, if married
with a normal man, will produce two normal visioned
female, one normal visioned male and one color blind male.
Cross 2
When a color blind female marry a normal visioned male

XCXC x XY………….Parents

XC XC X Y……Gametes

XCX XC Y XCY…………………F1
XC X

XCX (carrier female )and XCY (COLOR BLIND MALE) are

formed IN f1 GENERATION.
Daughters are carrier and son is colorblind.
When F1 daughter marries with colorblind male

F1 carrier daughter x Colorblind male

XCX XCY

XC X XC Y

XCXC XCY XCX XY……F2

Color blind daughter color bind son carrier daughter normal son
when a color blind woman is married with a normal visioned
man, in F1 generation they produce all color blind sons and
normal daughters which are carrier too.

When these carrier daughter marries with colorblind male, the

colorblind sons and daughter are produced in equal number.

Probable questions for exam?

1. What is criss-cross inheritance?
1. Describe sex linked inheritance with reference to eye color in
Drosophila.
2. Describe sex linked inheritance with reference to
colorblindness in man.
HW: Significance of criss-cross inheritance from book.
Crossing over
Crossing over is the exchange of chromosomal
segment between non-sister chromatids of
homologous chromosomes.

• The chromatids formed after crossing over are

called recombinants.

• As a result of crossing over new combinations of

DNA are formed.
• Crossing over takes place Pachytene stage of
Prophase I of meiosis I of cell division.
Mechanism of crossing over
It can be described in following steps.
1. Synapsis
The paternal and maternal chromosomes of
homologous pair come closed to each other at the
zygotene stage of prophase-I .

The process is called synapsis and homologous

chromosomes are called bivalent.

2. Duplication of chromosome
• Synapsis is followed by duplication of chromosome .

During diplotene stage the homologous

chromosome in bivalent splits longitudinally into
two sister chromatid forming a tetrad.
3. Crossing Over

The paired homologous chromosome start separating

at pachytene stage except one or more point known
as chaisma.

At this point, the non sister chromatid exchange the

genetic materials. This process is crossing over.

4. Terminalization

After the completion of crossing over the non sister

chromatid repel each other during diakinesis stage.

The chaisma shifts towards the terminal end known as

Terminalization.
Fig: Mechanism of crossing over
 Types of crossing over

1. Single crossing over

2.Double crossing over

3.Multiple crossing over

(HW:definition from book)
Significance of crossing over

1. It produces new combination of genes.

2. Useful recombinants can be formed by plant
breeders.
3. Various organism have been created as a
result of crossing over.
4. It helps in development of hybrid vigour.
5. It creates raw material for evolution.
6. It creates organism with better adaptability
and resistivity.