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Principles of Plant Genetics and Breeding

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the same plant. However, the dual color is impossible<br />

to reproduce by either sexual or asexual propagation.<br />

Commercial use <strong>of</strong> chimera is not attractive because the<br />

vegetative propagules must, <strong>of</strong> necessity, comprise both<br />

kinds <strong>of</strong> tissues in order to reproduce the maternal<br />

features.<br />

Gene action: dominant versus recessive mutations<br />

As previously indicated, mutations may cause a dominant<br />

allele to be changed into a recessive allele (recessive<br />

mutation), or a recessive allele to be changed into a<br />

dominant allele (dominant mutation). Open-pollinated<br />

species may accumulate a large amount <strong>of</strong> recessive<br />

mutant alleles without any adverse effects. However,<br />

upon selfing, the recessive alleles become homozygous<br />

<strong>and</strong> are expressed, leading to the phenomenon <strong>of</strong><br />

inbreeding depression. Using recessive genes in breeding<br />

takes a longer time because it requires an additional<br />

step <strong>of</strong> selfing in order to identify <strong>and</strong> select the desired<br />

recombinants. On the other h<strong>and</strong>, dominant mutations<br />

manifest in the current generation, needing no additional<br />

regeneration to be observable.<br />

Structural changes at the chromosomal level<br />

Three types <strong>of</strong> structural changes in the chromosome<br />

can occur as a result <strong>of</strong> mutation.<br />

Gene mutation<br />

Kind <strong>of</strong> mutation Gene mutations entail a change in<br />

the nucleotide constitution <strong>of</strong> the DNA sequence,<br />

adding or deleting nucleotides.<br />

1 Transitions <strong>and</strong> transversions. As previously<br />

described, the DNA consists <strong>of</strong> four bases – A, T, C,<br />

<strong>and</strong> G – that pair in a specific pattern, G–C <strong>and</strong> A–T.<br />

G<br />

C<br />

(a)<br />

G<br />

C<br />

A<br />

C<br />

Normal<br />

A<br />

T<br />

G<br />

C<br />

MUTAGENESIS IN PLANT BREEDING 201<br />

Mutant<br />

Normal<br />

Transition<br />

Transversion<br />

A T A C<br />

A T C G<br />

T A G C<br />

Figure 12.1 Mutations may occur by transition or<br />

transversion.<br />

The structure <strong>of</strong> the DNA may be modified in two<br />

ways – transition or transversion <strong>of</strong> the bases (Figure<br />

12.1). Mutation by transition entails the conversion<br />

<strong>of</strong> one purine base to another purine (or a pyrimidine<br />

to another pyrimidine). During replication, the second<br />

purine (a different purine), which has altered<br />

base-pairing properties, guides an incorrect base into<br />

position. Consequently, one normal base pair is converted<br />

to another pair that is genetically incorrect.<br />

An agent <strong>of</strong> mutation (a mutagen) such as nitrous<br />

acid has been known to cause deamination <strong>of</strong> adenine<br />

to hypoxanthin, cytosine to uracil, <strong>and</strong> guanine to<br />

xanthine, the net effect being a replacement <strong>of</strong> A–T<br />

with G–C in the DNA structure. A transversion<br />

involves the substitution <strong>of</strong> a purine by a pyrimidine<br />

<strong>and</strong> vice versa.<br />

2 Tautomeric shifts. It is known that each <strong>of</strong> the bases<br />

<strong>of</strong> DNA can exist in rare states as a result <strong>of</strong> the redistribution<br />

<strong>of</strong> electrons <strong>and</strong> protons in the molecule,<br />

events called tautomeric shifts. When this occurs, the<br />

base sometimes is unable to hydrogen bond with its<br />

complementary base. Instead, some <strong>of</strong> these altered<br />

bases succeed in bonding with the wrong bases,<br />

resulting in mutations when, during replication, one<br />

purine (or pyrimidine) is substituted for the other<br />

(Figure 12.2).<br />

3 Effect <strong>of</strong> base analogues. Certain analogues <strong>of</strong> the<br />

naturally occurring bases in the DNA molecule have<br />

Figure 12.2 Mutations may be caused by tautomeric shifts: (a) shift involving cytosine, <strong>and</strong> (b) shift involving thymine.<br />

A<br />

T<br />

(b)<br />

A<br />

T<br />

G<br />

T<br />

Normal<br />

G<br />

C<br />

A<br />

T<br />

Mutant<br />

Normal

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