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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.
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 (%)
X 100
Total number of progenies
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.