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LINKAGE

The tendency of two or more genes to stay together during inheritance is


known as linkage. It is the consequence of the concerned genes being located in
the same chromosome.
Linked genes do not show independent segregation; as a consequence, the
ratios obtained in F2 and test-cross generations are significantly different from
the expected ratios of 9:3:3:1 and 1:1:1:1 (in two linked genes).
Experiment on Pea
Bateson and Punnet in 1905 studied the inheritance of flower colour and
pollen shape in pea. A dominant gene B produces blue flowers, while its
recessive allele b determines red flowers. Another dominant gene L governs
elongate pollen grains, whereas its recessive allele l is responsible for normal
round pollen grains. When the F1 from the cross blue elongate (BB LL) x red
round (bb ll) was test crossed (with red round, bb ll), the progeny showed the
ratio 7 blue elongate (Bb L1) : 1 blue round (Bb ll) : 1 red elongate (bb Ll) : 7 red
round (bb ll) in place of the expected 1 : 1 : 1 : 1 ratio. It appears as if the two
dominant genes (B and L) have an affinity for each other so that they tend to
stay together during inheritance; Bateson and Punnet called this situation as
coupling phase. But when the F1 from the cross blue round (BB ll) x red
elongate (bb LL) was test crossed (with red round, bb ll), the progeny exhibited
the ratio 1 blue elongate (Bb Ll) : 7 blue round (Bb ll) : 7 red elongate (bb Ll) : 1
red round (bb ll). It appears as if, in this case, the two dominant genes (B and
L) repel each other so that they tend to stay away from each other during
inheritance. This situation was referred to as repulsion phase.
Bateson and Punnet failed to advance a suitable explanation for these
findings. They suggested that there was a selective multiplication (through
mitosis) of some types of gametes, viz., of BL and bl in the coupling phase, and
of Bl and bL in the repulsion phase, after meiosis; this gave rise to the 7 : 1 : 1 :
7 and 1 : 7 : 7 : 1 testcross ratios in place of the expected 1 : 1 : 1 : 1. This
hypothesis is untenable as it is contrary to the known facts. But the terms
coupling phase and repulsion phase are widely used to denote linkages between
two (or more) dominant genes, and between one (or more) dominant and one (or
more) recessive genes, respectively.
Experiment on Maize
In maize, a dominant gene C produces coloured seeds, while its
recessive allele, c determines colourless seeds. Another dominant gene Sh
governs full seeds, whereas its recessive allele sh gives rise to shrunken seeds.

When plants having coloured full seeds (CC ShSh) were crossed with the having
colourless shrunken seeds (ccshsh), F1 seeds were coloured full (Cc Shsh). Out
of the 8,368 seeds obtained from the test cross Cc Shsh X
ccshsh, 4,032
(48.2%) were coloured full 4035 (48.3%) were colourless shrunken, 149 (1.7%)
were coloured shrunken and 152 (1.8%) were colourless full. Clearly, the four
phenotypic classes are not presented in the expected ratio of 1:1:1:1. The
phenotypic classes of the parental combinations are much higher than the
expected. They are known as parental phenotypes (coloured full and colourless
shrunken). The remaining two phenotypic classes, coloured shrunken and
colourless full are far less frequent than expected (25%). These two character
combinations are called recombinant phenotypes, since they are generated by
reshuffling of the characters of the two parents of the F1 used in the test cross.

Fig. Coupling Phase Linkage in Maize

In the above example, it appears as if the two dominant gene C and Sh


have a strong affinity for each other so that the frequencies of coloured full and
colourless shrunken phenotypes are greater than expected. This situation is
referred to a coupling phase, (Fig 14.1) and is due to the presence of genes C
and Sh in the chromosome.
Similarly, when plants having coloured shrunken seeds (CC shsh) were
crossed with those having colourless full seeds (ccShSh), the F 1 seeds were
coloured full
(CcShsh). But when F 1 plants were test-crossed (CcShsh)
x (ccshsh), 47.9 % of the seeds were coloured shrunken, 49.1 % were colourless
full, 1.4 % were coloured full and

Fig. Repulsion Phase Linkage in Maize

1.5 % were colourless shrunken. In this case also, the parental types (colourless
full and coloured shrunken) were more frequent while the recombinant types
(coloured full and colourless shrunken) were less frequent than expected. It

appears as if the dominant genes C and Sh dislike each other so that the
phenotypic classes coloured shrunken and colourless full are much more
frequent than expected. This situation is referred to as repulsion phase (Fig
14.2); it is due to the presence of the dominant allele of one gene, e.g., C, with
the recessive allele of the other gene, e.g., sh, in the same chromosome.
Obviously, coupling and repulsion phases are only the two situation of linkage.
Incomplete Linkage
When only parental character combinations, are recovered in the test
cross progeny, it is called complete linkage. Sometimes, their alleles recombine
to produce recombinant gametes, e.g., Csh and cSh when such recombinant
(due to crossing over) types are also recovered, in addition to the two parental
types, in the test cross progeny, it is called incomplete linkage.
CROSSING OVER
The exchange of precisely homologous segments between non-sister
chromatids of homologous chromosomes is known as crossing over. It is
responsible for recombination between linked genes. Crossing over takes place
during pachytene. During pachytene, each chromosome of a bivalent
(chromosome pair) has two chromatids so that each bivalent has four
chromatids or strands (four stranded stage).
Generally, one chromatid from each of the two homologous of a bivalent
is involved in crossing over
(Fig ). The point of exchange of the homologous
segment of the non-sister chromatids of homologous chromosome is known as
chiasma. Obviously, each event of cross over produces two recombined
chromatids, called crossover chromatids, and two original chromatids referred
to as non-crossover chromatids.
The frequency of crossing over between two genes can be estimated as
the frequency of recombinant progeny from a test cross for these genes. It is
expressed in percentage.

Frequency of crossing =
over (%)

Number of recombinant progenies


from a test cross

X 100
Total number of progenies

Fig. Crossing Over Between Non-sister Chromatids

Crossing over in four-strand stage (Experiments on Neurospora)


In Neurospora, each zygote undergoes meiosis to yield four haploid
nuclei; these nuclei undergo mitosis to yield eight haploid (n) nuclei. They are
arranged in a linear order; each nucleus gives rise to one ascospore (Fig ).
The dominant gene Al of Neurospora produces black ascospores, while
its recessive allele al determines albino ascospores. The regular arrangement of
ascospores makes it possible to predict the consequence of crossing over
between this gene and its centromere on the spore arrangement.
If there is no crossing over between the gene and the centromere or if
crossing over takes place between two-strand stage, the ascospores obtained
from the heterozygous Alal, zygotes would show 4 black: 4 albino spore
arrangement. However, if the crossing over occurred in four strand stage, the
spores would show a 2 black : 2 albino : 2 black : 2 albino or 2 :4:2
arrangement.
The asci from Alal heterozygotes show 4 : 4, 2 : 4 : 2 as well as 2 : 2 :
2 : 2 spore arrangement. The 4 : 4 arrangement may be due to no crossing over
or due to crossing over in two-strand stage. But 2 : 4 : 2 and 2 : 2 : 2 : 2
arrangement in some of the asci of Alal heterozygotes clearly show that
crossing over takes place in four-strand stage.

Crossing Over between three linked genes

Fig. Crossing over between three linked genes

A test cross for three linked genes eg., c, sh and wx in maize, yields
eight
types
of
gametes
and
phenotypes
(Fig.). Two of these types i.e., CWxSh and cwxsh are the most frequent and
represent the parental or non-recombinant types, while two others i.e., CwxSh
and cWxsh are the least frequent and are double cross overs. The remaining
four types i.e., Cwxsh, cWxSh, cwxSh and CWxsh are produced by single
cross overs between three linked genes, and the frequency is intermediate
between the parental and double cross over types.

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