5/21/2021

Genetic linkage,
Recombination & mapping
Dr. Fariha Khan
BIO231
BSC208

Independent assortment:
Two genes on two different homologous pairs of chromosomes

Two genes on nonhomologous

chromosomes (unlinked genes)
yield 4 kinds of gametes in equal
proportions, 25% frequency

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Linkage: Two genes on a single pair of

homologs; no exchange occurs

➢ If no crossing over occurs between

the two genes , only two
genetically different gametes are
formed.
➢ Each gamete receives the alleles
present on one homolog or the
other, which is transmitted intact as
the result of segregation.

➢ This case demonstrates complete

linkage, which produces only
parental or noncrossover gametes

50% AB; 50% ab

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Linkage and Recombination

• Because the chromosome, not the gene, is the unit of transmission
during meiosis, linked genes are not free to undergo independent
assortment.
• Instead, the alleles at all loci of one chromosome should, in theory, be
transmitted as a unit during gamete formation.
• However, in many instances this does not occur because of crossing
over event during the first meiotic prophase
• This crossing over event results in the reshuffling, or recombination,
of the alleles between homologs
• Thus alleles of linked genes can get separated by crossing over
(homologous recombination).

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Figure shows the results of crossing over

between two linked genes. As you can
see, this crossover involves only two
nonsister chromatids of the four
chromatids present in the tetrad. This
exchange generates two new allele
combinations, called recombinant or
crossover gametes. The two chromatids
not involved in the exchange result in
nonrecombinant gametes

When genes are far apart, crossing

over happens often enough that all
types of gametes are produced with
25% frequency

When genes are very close together on the

same chromosome, crossing over still
occurs, but the outcome (in terms of
gamete types produced) is different.
Instead of assorting independently, the
genes tend to "stick together" during
meiosis.
That is, the alleles of the genes that are
already together on a chromosome will
tend to be passed as a unit to gametes. In
this case, the genes are linked.
Now, we see gamete types that are present
in very unequal proportions

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Linkage and Recombination

▪ The closer together two genes are on a chromosome, the less likely their alleles will be
separated by crossing-over.
▪ Genes that are farther away from each other are more likely to be separated during a
process of homologous recombination
▪ Genes on separate chromosomes are not linked, and assort independently
▪ The frequency of crossing over between any two loci on a single chromosome is
proportional to the distance between them.

Key points:
➢When genes are found on different chromosomes or far apart on the same
chromosome, they assort independently and are said to be unlinked.
➢When genes are close together on the same chromosome, they are said to be
linked. That means the alleles, or gene versions, already together on one
chromosome will be inherited as a unit more frequently than not.
➢Linkage between genes is detected as a deviation from expectations based on
Mendel’s Principle of Independent Assortment.
➢Linkage between genes causes them to be inherited together and reduces
recombination; crossing over breaks up the associations of such genes.

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Recombination frequency
• The frequency with which crossing over occurs between any two
linked genes is generally proportional to the distance separating the
respective loci along the chromosome.
• The percentage of recombinant progeny produced in a cross is called
the recombination frequency, which is calculated as follows:

(r)

Linkage between the

genes for flower color and P
pollen length in sweet
peas.
F1

Nonrecombinant Recombinant
F1 gametes chromosomes chromosomes
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Testcross for linkage between

genes in sweet peas:
F1

Because the recombinant progeny

in the F2 are 8 percent of the total,
the genes for flower color and
pollen length are tightly linked.

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Recombination Frequency
Recombinant = number of recombinants produced 100%
Frequency Total number of gametes

= genetic Map distance= Map unit (m.u)= centiMorgen (cM)

• Two genes that have recombination frequency of 50% are located

on nonhomologous chromosome or far apart on same
chromosome= unlinked genes

• Genes with recombination frequency less than 50% are on same

chromosome= linked genes

• The frequency of recombination measures the intensity of

linkage. In the absence of linkage, this frequency is 50%; for
very tight linkage, it is close to zero
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Recombination Frequency and Gene/chr. Mapping

Linked genes can be mapped on a chromosome by studying how often
their alleles recombine
➢ We can see if two genes are linked, and
how tightly, by using data from genetic
crosses to calculate the recombination
frequency.
➢ By finding recombination frequencies for
many gene pairs, we can make linkage
maps that show the order and relative
distances of the genes on the
chromosome.
The correlation of RF allows us to construct chromosome
maps, which give the relative locations of genes on
chromosomes.
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Recombination Frequency and Genetic Distance

➢The frequency with which the genes are unlinked by
crossovers is directly proportional to how far apart they
are on their chromosome.
➢The recombination frequency gives a measure of the
distance between two genes or loci
➢If recombination frequencies for different pairs of genes
are estimated, a map of relative positions of genes on
the chromosome can be constructed.
➢Two loci that show 1% recombination are defined as
being 1 centiMorgan apart on a genetic map.
1 genetic map unit =1cM (centiMorgan)

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Example : Genetic Map Construction

• In a series of experiments investigating three characteristics A, B
and C it was found that:
• Recombinant frequency b/w A and B = 1.0%.
• Recombinant frequency b/w A and C = 0.6%
• Recombinant frequency b/w B and C = 0.4%.

A genetic map of the three genes of these characteristics A, B, C :

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For example:
let's suppose we have three genes, A, B, and C, and we want to
know their order on the chromosome (ABC? ACB? CAB?)

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Question
• Genes A, B, C, and D are located on the same chromosome. The
recombination frequencies (RF) are as follows:
Relationship RF
A-B 10%
A-C 25%
A-D 23%
B-C 15%
C-D 48%
What is the most likely order of the genes on the chromosome?
CBAD

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