Genetic Linkage

Homologous chromosomes

A chromosome is a single piece of DNA. Genes are segments of DNA arranged along a chromosome. A
single chromosome can have hundreds or even thousands of genes. The specific position where a
particular gene is located on a chromosome is called a locus (loci). Chromosomes exist in pairs in diploid
organisms, with each pair containing one chromosome from each parent. Homologous chromosomes
are a pair of chromosomes that have the same genes arranged in the same order but may have different
alleles for those genes. Alleles are different versions of the same gene. Homologous chromosomes are
similar in length, centromere position, and banding pattern, making them structurally alike.

Recombination

Recombination, also known as crossing over, is a process that occurs during the pachytene stage of the
prophase of meiosis 1. It involves the exchange of genetic material between homologous chromosomes
at points called chiasmata. This exchange occurs when non-sister chromatids of homologous
chromosomes break and rejoin, swapping equivalent segments of DNA. Recombination is crucial for
increasing genetic diversity in sexually reproducing organisms by creating new combinations of alleles.

Characteristics of Recombination

-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.

-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.

Linkage

Linkage is the phenomenon in which certain genes stay together during inheritance through generations
without any change or separation due to their presence on the same chromosome. Linkage was first
suggested by Sutton and Boveri (1902-1903) when they propounded the chromosomal theory of
inheritance, which states that genes are located on chromosomes, and the behavior of chromosomes
during meiosis explains how genetic traits are passed from parents to offspring. However, in 1910
Thomas Hunt Morgan clearly proved and defined linkage on the basis of his breeding experiments in
Drosophila melanogaster. In 1911, Morgan and Castle proposed chromosome theory of linkage. It states
that;

(i) Linked genes occur in the same chromosome.

(ii) They lie in a linear sequence in the chromosome.
(iii) There is a tendency to maintain the parental combination of genes except for occasional
crossovers.
(iv) Strength of the linkage between two genes is inversely proportional to the distance between
the two, (i.e., two linked genes show higher frequency of crossing over if the distance
between them is higher and lower frequency if the distance is small).

Unlinked genes show independent assortment, a dihybrid ratio of 9: 3: 3: 1 and the di-hybrid or
double test cross ratio of 1: 1: 1: 1 with two parental and two recombinant types. Linked genes do
not show independent assortment but remain together, producing only parental type of progeny.
They give a di-hybrid ratio of 3: 1 and a test cross ratio of 1: 1.

Types of Linkage:

Complete Linkage: Complete linkage occurs when two genes are located very close to each other on the
same chromosome, such that they are almost always inherited together without any recombination. In
this case, the parental combination of alleles is maintained across generations, and no new gene
combinations arise. This phenomenon results in gametes that exclusively contain parental genotypes.

Incomplete Linkage: Incomplete linkage occurs when two genes are located on the same chromosome
but are far enough apart that recombination can occur between them. This leads to the production of
both parental and recombinant gametes, resulting in a mixture of parental and new gene combinations
in the offspring. Incomplete linkage is more common than complete linkage and contributes to genetic
variation in populations.

Linkage Groups: A linkage group is a linearly arranged group of linked genes which are normally
inherited together except for crossing over. It corresponds to a chromosome which bears a linear
sequence of genes linked and inherited together. Because the two homologous chromosomes possess
either similar or allelic genes on the same loci, they constitute the same linkage group. Therefore, the
number of linkage groups present in an individual corresponds to number of chromosomes in its one
genome (all the chromosomes if haploid or homologous pairs if diploid).
Recombination Frequency:

Recombination frequency is a measure of the likelihood that a crossover event will occur between two
genes during meiosis, resulting in the exchange of genetic material. It is a calculation to define the
number of parental and recombinant gametes. Parental genotypes are the combinations of alleles in
offspring that are identical to those found in the parents, resulting from genes being inherited without
any recombination. Recombinant genotypes are new combinations of alleles in offspring that arise due
to recombination (crossing over) between homologous chromosomes during meiosis.

The equation is as follows:

Recombination Frequency = The number of recombinant offspring / total number of offspring x 100%

The recombination frequency cannot be higher than 0.50. If alleles assort independently, there will be a
random distribution of alleles in the progeny — 50% will be recombinant gametes and 50% will be
parental gametes, making the RF approximately 0.50. If a gene is linked, there will be a higher
percentage of parental gametes, making the RF < 0.50.