1. Genetic linkage occurs when two genes located near each other on the same chromosome tend to be inherited together during meiosis. 2. Early theories of linkage proposed by Sutton, Boveri, Bateson and Punnett failed to fully explain observed inheritance patterns. 3. Morgan's chromosomal theory of linkage established that genes are linearly arranged on chromosomes and that the closer two genes are, the stronger the tendency for them to be inherited together. This provided an explanation for linkage patterns and laid the foundation for modern genetics.

1. Linkage – Concept, Types and
theories
Vaishali S.Patil
Professor, Department of Botany
Shri Shivaji College of Arts, Commerce & Science
Akola

2. Linkage-
Genetic linkage describes the way in which two genes that are
located close to each other on a chromosome are often inherited
together, during the meiosis phase of sexual reproduction.
1.Each chromosome contains more than one gene.
2.The genes for different characters may be either situated in the
same chromosome or in different chromosomes.
3.When the genes are situated in different chromosomes, the
characters they control appear in the next generation together or apart,
depending on the chance alone. They assort independently according
to Mendel’s law of independent assortment.
4.But if the genes are situated in the same chromosome and are fairly
close to each other, they tend to be inherited together.
This type of coexistence of two or more genes in the same
chromosome is known as linkage.

4. The occurrence of different genes on the same chromosome. They show non- random
assortment at meiosis.
Characteristic of Linked Genes
Genes on different chromosomes assort independently giving a 1: 1: 1: 1 test cross ratio.
Linked genes do not assort independently but tend to stay together in the same
combination as they were in the parents.
Types of Linkage-
Morgan and his co-workers by their investigation on the Drosophila found two types of
linkage –
1.Complete linkage and
2.Incomplete linkage.
Complete Linkage-
The complete linkage is the phenomenon in which parental combinations of characters
appear together for two or more generations in a continuous and regular fashion.
In this type of linkage genes are closely associated and tend to transmit together. These
genes do not conform to Mendel’s principle of independent assortment they fail to
produce the expected 9:3:3:1 ratio in a breeding situation involving the inheritance of
two pairs of contrasting characters (Dihybrid cross).

6. Complete Linkage:
If linkage is complete, there should be parental combinations only and
no recombination. Morgan (1919) reported a complete linkage in
Drosophila. When ordinary male wild fly with grey body and normal
wings was crossed with female having black body and vestigial wings, in
F1, hybrids were all grey bodied and normal winged (with dominant
characters).
But when F1, male is back crossed with recessive female parent only two
types of individuals in F2 generation were produced instead of expected
four. These two types were grey bodied and normal winged and black
bodied and vestigial winged in equal number, thus indicating the
complete linkage.
But how do genes located on same chromosome assort independently? It
may be due to exceptions in Mendel’s second law of independent
assortment or due to some mechanism for genes on the same
chromosome to separate and recombine during meiosis. Both the
cases have been observed.

7. Incomplete linkage:
Incomplete linkage produces new combinations of the genes in the
progeny due to the formation of chiasma or crossing over in between
the linked genes present on homologous chromosomes. When in sweet
peas a cross is made between blue flower and long pollen (BBLL) with
red flower and round pollen (bbll) in F1 expected blue flower and long
pollen (BbLl) heterozygous condition is got. (Fig. 5.47).
However, test cross between blue and long (BbLl) and double
recessive (bbll) gave blue long (43.7%), red round (43.7%), blue
round (6.3%) and red long (6.3%). The parent combinations are
87.4% are due to linkage in genes on two homologous chromosomes,
while in case of new combinations (12.6%) the genes get separated
due to breaking of chromosomes at the time of crossing over in
prophase-I of meiosis. New combinations in the progeny appeared
due to incomplete linkage (Fig. 5.47).

9. Significance of Linkage:
1. Linkage does not permit the breeders to bring the desirable
characters in one variety.
2. Linked characters are maintained for generations because
linkage prevents the incidence of recombination.
Thus linkage has one of the great significance for the living
organisms in that it reduces the possibility of variability in
gametes unless crossing over occurs.

10. Theories-1. W. Sutton and T. Boveri 1902) 2.Sutton (1903) and Bateson and Punnett (1906)3. T.
H Morgan (1911)
1.W. Sutton and T. Boveri 1902
Theory-
On the Boveri–Sutton chromosome theory is a fundamental theory of genetics which
identifies chromosomes as the carriers of genetic material. It correctly explains the
mechanism underlying Mendels laws of inheritance by identifying chromosomes with
the paired factors required by Mendels laws. It also States that chromosomes are
composed of linear structures of genes located at specific sites called loci along them.
It States simply that chromosomes, which can be seen in all dividing cells and pass
from one generation to another, is the basis for all genetic inheritance. Over time
random mutations involve changes in the DNA sequence of the gene. Genes are located
on chromosomes.

11. The chromosome theory of inheritance is credited to papers by
Walter Sutton in 1902 and 1903, as well as to independent work by
Theodor Boveri during roughly the same period. Boveri was studying
sea urchins, in which he found that all chromosomes must be present for
normal embryonic development to take place. Suttons work with
grasshoppers showed that chromosomes occur in matched pairs of
maternal and paternal chromosomes which separate during meiosis
and "may pose a physical basis of the law of heredity Mendel".

14. 2.Coupling and Repulsion hypothesis of linkage: by
Bateson and Punnett
•Bateson and Punnett in 1906 described a cross in sweat pea (Latharus odoratus),
where they find gene pairs do not assort independently as expected.
•While they were working on sweet pea, they noticed two variety of plant-plant with
Blue flowers (BB) with long pollen grains (LL) and red flower (bb) with round pollen
grain (ll).
•When the sweat pea variety having blue flower and long pollen grain (BBLL) were
crossed with those of another variety having red flower and round pollen (bbll).
In F1 generation, all the offspring having blue flower and long pollen (BbLl) were
observed as expected. Because, blue color is dominant over red and long pollen is
dominant over round.
•When these F1 offspring (BbLl) were test crossed with red flower and round pollen
variety (bbll), they obtained (BbLl), (Bbll), (bbLl), (bbll) in the ration 7:1:1:7 instead of
1:1:1:1.
•This indicates that the dominant characters (Blue color and long pollen) tends to
remain together. Similarly, the recessive characters (red color and round pollen) tends to
remain together. This deviation or phenomenon is explained by Coupling and
Repulsion hypothesis by Bateson and Punnett.

15. •Bateson and punnett conclude that- the pairs of genes from homozygous parents try to
remains together in a cross due to which parental characters is more frequent. This is
due to coupling.
•Two dominant gene for Blue flower and long pollen (BL) are linked in one plant while
in other plant two recessive gene for red flower and round pollen (bl) are linked. Such
linkage is called coupling.
• In another experiment, when a cross was made between sweet pea variety having blue
flower and round pollen (BBll) with another variety having red flower and long pollen
(bbLL)
•In F1 generation, all offspring with blue flower and long pollen were obtained (BbLl).
When these F1 offspring (BbLl) were test crossed with red flower and round pollen
variety (bbll), they obtained (BbLl), (Bbll), (bbLl), (bbll) in the ratio 1:7:7:1 instead of
1:1:1:1.
•It was observed in both experimental cases that plants with parental characters are
obtained in excess number than the plants with recombinant characters. Also
when dominant alleles are from different parents (heterozygous ) the frequency of
recombination is more because of repulsion of gene (repulsion).
In First experiment, BL are linked and such linkage is known as Coupling and in
second experiment bL are linked and such linkage is known as Repulsion.
The coupling and repulsion hypothesis is now discarded.

16. Chromosomal theory of linkage: by T. H Morgan
•Bateson and Punnett failed to explain the exact reasons of coupling and repulsion
•Later, T.H. Morgan who found coupling and repulsion hypothesis incomplete, while
performing experiments with Drosophila, in 1910.
•He proposed that the two genes are found in coupling phase because they are present
on same chromosome and similarly on repulsion phase because they are preset on
two different homologous chromosomes.
• There genes are then called linked genes and the phenomenon of inheritance of such
linked genes is called linkage by Morgan.
• The term coupling and repulsion were replaced by the terms, cis and trans by
(Haldane, 1942).
•Morgan stated the linked genes have the tendency to remain together in original
combination because they are located on same chromosome. And the strength of
linkage depends upon the distance between the linked genes in the chromosome.
The concept of linkage by Morgan establish the foundation of Cytogenetics and develop
the theory of linear arrangement of genes in the chromosomes and helps to construct
genetic map of the chromosome.

17. According to Chromosomal theory of linkage:
•Chromosome contains genes.
•Genes lie in a linear order in a chromosome
• Distance between them is variable.
•Each gene has a definite locus in a chromosome.
•The genes which are close to each other, shows the phenomenon of
linkage.
•The linked genes cannot be separated during gametogenesis (inheritance
process), they inherited together.
•Tendency of genes to remain linked is due to their presence on same
chromosome.
•The distance between the linked genes determines the strength of
linkage.
•The closer the distance stronger is the linkage strength.
•The linkage is not due to any relation between two genes but is simply
because they happens to be located in the same chromosome.