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Mechanisms and Inheritance of Resistance to Insect Pests 131<br />

Escape<br />

Avoidance of insect damage through early or late fl owering during periods of low insect<br />

abundance is another mechanism of resistance, called pseudo-resistance (Kogan, 1982;<br />

Smith, 2005). It is not a true mechanism of resistance, but serves as an effective means of<br />

avoiding pest damage. Many of the landrace cultivars have evolved such that they fl ower<br />

during the periods of low insect activity. This mechanism can be exploited to minimize<br />

insect damage, depending on the length of the cropping season, and the cultivars grown.<br />

Breeding for Resistance to Insect Pests<br />

Several methods have been used to transfer insect resistance genes into high-yielding cultivars<br />

(Panda and Khush, 1995; Smith 2005). Mass selection, pure line selection, recurrent<br />

selection, and backcross methods have been used to incorporate insect resistance genes<br />

into the crop plants. In self-pollinated crops, backcross, bulk, and pedigree breeding have<br />

been used to incorporate resistance genes into agronomically desirable cultivars, while<br />

mass selection and recurrent selection methods have been used in cross pollinated crops.<br />

Mass Selection<br />

Mass selection involves selecting individual insect-resistant plants in a population in each<br />

generation (cycle of selection), combining the seed of the resistant plants, and growing<br />

progenies in the following generation. It allows selection of several insect-resistant plants<br />

in each cycle of selection. Maximum improvement for resistance is achieved during initial<br />

selection, followed by two to fi ve additional cycles of selection. Mass selection has been<br />

used effectively to breed for resistance to the potato leafhopper, Empoasca fabae (Harris)<br />

(Sanford and Ladd, 1983) and to identify genotypes with resistance to sugarbeet root maggot,<br />

Tetanops myopaeformis Roder (Theurer et al., 1982). Pure line selection is another form of<br />

mass selection that involves the selection of individual resistant plants that are advanced<br />

separately. In each cycle of selection, resistant selections are selfed and advanced. In crosspollinated<br />

crops, individual selections are interplanted in a later selection cycle to form a<br />

composite of all the insect-resistant plants/lines.<br />

Recurrent Selection<br />

Recurrent selection has been used to concentrate insect resistance genes from diverse<br />

sources into the same background through several cycles of intermating and selection. In<br />

each cycle, resistant plants are selected among the progeny produced through a previous<br />

mating of resistant individuals. Recurrent selection allows introduction of resistance to<br />

several insects and diseases from different sources in later selection cycles. Recurrent selection<br />

has been used to increase the levels of resistance in maize to European corn borer<br />

O. nubilalis (Russell, Lawrence, and Guthrie, 1979; Klenke, Russell, and Guthrie, 1986). The<br />

corn borer damage decreased signifi cantly after four cycles of selection. Variability for<br />

resistance to O. nubilalis also decreased in each cycle of selection. Recurrent selection has<br />

also been used to increase the level of resistance in maize populations to corn earworm,<br />

H. zea, and fall armyworm, Spodoptera frugiperda ( J.E. Smith) (Widstrom, 1989; Widstrom et al.,

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