Linkage

S J BORKAR
• The tendency of genes present on same chromosome to enter into
same gamete during process of gametogenesis is called Linkage.
• Genes on the same chromosome are like passengers on a charter
bus: they travel together and ultimately arrive at the same
destination.
• A few years after the rediscovery of Mendel’s laws of inheritance,
Bateson and Punnett (1905) observed in sweet pea that two
pairs of alleles do not assort independently. Morgan (1910) found
the same phenomenon in drosophila and Hutchinson observed a
clear cut case of linkage in maize.
• All these researches found that genes inherit in groups rather
than individually. This tendency of two or more genes to remain
together in the same chromosome during inheritance is referred to
as linkage. In other words, linkage is the tendency of genes to be
inherited in groups.
• This is a deviation from the Mendelian principle of
independent assortment.
• Mendel’s law of independent assortment is applicable to the genes
that are situated in separate chromosomes. When genes for
different characters are located in the same chromosome, they are
tied to one another and are said to be linked.
 Postulates of Theory of Linkage
• Linkage involves two or more genes which are located in the same chromosome in a linear fashion.
• Linkage may involve either dominant genes or recessive genes or some dominant and some recessive genes.
• Linkage usually involves those genes which are located closely.
• Presence of linkage leads to higher frequency of parental types than recombinants in a test cross progeny. When
two genes are linked the segregation ratio of a test cross progeny deviates significantly from the 1 : 1 : 1 : 1 ratio.
• Linkage may involve either two or more desirable traits or all undesirable traits or some desirable and some
undesirable traits.
• Linkage is observed for both oligogenic traits as well as polygenic traits. However, it is more common for the
former than latter.
• Besides pleiotropy, linkage is an important cause of genetic correlation between various plant characters.
• The strength of linkage depends on the distance between the linked genes. Lesser the distance higher the strength
and vice versa.
• If crossing over does not occur, all the genes located in one chromosome are expected to be inherited together.
Thus the maximum number of linkage groups in an organism is equal to its haploid chromosome number.
• Linkage can be broken by repeated inter-mating of randomly selected plants in segregating populations for
several generations.
Phases of Linkage of Genes:
• There are two phases of linkage, viz., coupling phase and repulsion phase. These phases
were given by Bateson and Punett (1905), but they could not give proper interpretation of
these terms.
• Later on, Morgan (1910) based on his studies with Drosophila explained that coupling and
repulsion are the two aspects of the same phenomenon what we call linkage. The coupling
and repulsion phases of linkage are briefly described below.

Coupling:
• The linkage between two or more either dominant (AB) or recessive (ab) alleles is referred
to as coupling. A good example of coupling was reported by Hutchinson in maize for the
genes governing colour of seed (coloured and colourless) and shape of seed (full and
shrunken).
• The coloured seed is governed by dominant gene (C) and full seed is also governed by
dominant gene (S). He made cross between plants having coloured full seeds (CCSS) and
colourless shrunken seeds (ccss). The F1 seeds were coloured full. When the F1 was test
crossed with double recessive parent the following results were obtained instead of 1 : 1 : 1 :
1 ratio.
• This indicates that parental combinations are
higher than recombination’s, indicating
presence of linkage. The parental
combinations occurred in 96.4% instead of
50% and re-combinations were 3.6% instead
of 50% in this case.

• So when, Dominant alleles of linked genes

are present on one chromosome and
recessive alleles on homologous
chromosome. During gametogenesis
dominant alleles enter in same gamete
similarly recessive alleles also.
• This phase of linkage is called Coupling
Phase.
Repulsion:
• The linkage of dominant allele with that of the
recessive allele (Ab or aB) is known as repulsion.
Hutchinson also observed repulsion phase of
linkage in maize.
• He observed this type of linkage when he made
cross between plants having coloured shrunken
seeds (Cs) with those having colourless full seeds
(cS).
• In F1 the seeds were coloured full. By crossing of
F1 with double recessive parent the following
results were obtained instead of 1 : 1 : 1 : 1 ratio.

• So when, Dominant alleles of a gene is present

along the recessive allele of a linked gene on the
same chromosome.
• During gametogenesis dominant and recessive
alleles enter into different gametes.
• This phase of linkage is called Repulsion
Phase.
 Types of Linkage:
Complete Linkage
• Linkage in which crossing over
does not occur is known as
complete linkage or absolute
linkage.
• In other words, when only
parental types are obtained
from the test cross progeny, it
refers to complete linkage.
Good example of complete
linkage is Drosophila male and
female silk moth.
Incomplete Linkage

•The linked genes which are widely

located in chromosomes and have
chances of separation by crossing over
are called incompletely linked genes
and the phenomenon of their
inheritance is called incomplete
linkage.

•The linked genes do not always stay

together because homologous non-
sister chromatids may exchange
segments of varying length with one
another during meiotic prophase.

•This sort of exchange of chromosomal

segments in between homologous
chromosomes is known as crossing
over.
 Linkage Significance in Plant Breeding:
(i) Effect on Selection:
• Linkage between two or more loci controlling different desirable character is advantageous for a plant
breeder. Because desirable alleles will occur together more frequently in segregating population than would
be expected with independent assortment.
• Thus the breeder can easily obtain individuals with desirable alleles for two characters. Linkage is
undesirable when desirable and undesirable genes are linked together. To obtain sergeant with desirable
alleles, breeder has to grow larger populations than with unlinked loci.
(ii) Effect on Genetic Variance:
• The estimates of genetic variances for quantitative characters are greatly influenced by the presence of
linkage. Linkage disequilibrium influences the gene action by way of causing bias in the estimates of genetic
variances. Linkage in repulsion phase in segregating population causes an upward bias in the estimates of
dominance variance and downward bias in the estimates of additive genetic variance.
(iii) Effect on Genetic Correlation:
• Linkage has great impact on the genetic correlation. The linked characters generally show high values of
genetic correlation and also of co-heritability. A linkage between genes controlling two different desirable
characters will help in simultaneous improvement of both characters. If there is no linkage between two
desirable characters, the breeder has to combine such characters from two different sources.