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Mechanisms and Inheritance of Resistance to Insect Pests 133<br />

resistance in maize (G.E. Scott, Dicke, and Pesho, 1966; Onukogu et al., 1978). For this<br />

reason, maize hybrid development has been conducted using other breeding methods.<br />

Development of F1 Hybrids Using Cytoplasmic Male Sterility<br />

Male-sterility results from the inability of plants to produce functional pollen due to reproductive<br />

defi ciency in hermaphrodite fl owers. Cytoplasmic male-sterility occurs due to the<br />

mutation of mitochondria or some other cytoplasmic factors outside the nucleus, which<br />

results in the transformation of fertile cytoplasm into sterile cytoplasm. The male-sterile<br />

line is maintained by pollinating it with pollen from the maintainer line, which differs<br />

from the A line only for male-sterility. The F1 hybrids for cultivation are produced by<br />

crossing the male-sterile line with a pollinator that carries the fertility restoration genes.<br />

A pollinator line that results in maximum heterosis for grain yield and other traits of interest<br />

is used for hybrid production. Cytoplasmic male-sterility has been exploited in several<br />

crops to develop hybrids for increasing crop productivity, particularly in cereal crops such<br />

as rice, maize, sorghum, and pearl millet. However, large-scale cultivation of hybrids based<br />

on a single source of male sterility may pose a serious challenge to sustainable crop production<br />

because of decreased genetic diversity and increased susceptibi lity to insect pests.<br />

Therefore, there is a need to develop male-sterile and restorer lines with resistance to<br />

insects to develop insect-resistant hybrids for cultivation by the farmers. To produce insectresistant<br />

hybrids involving a cytoplasmic male-sterility system, it is important to transfer<br />

insect resistance genes into both male-sterile and restorer lines to produce hybrids with<br />

resistance to insects. Sorghum hybrids with resistance to shoot fl y and sorghum midge<br />

have been developed based on cytoplasmic male sterility (H.C. Sharma et al., 1996, 2005c;<br />

Dhillon et al., 2006a; H.C. Sharma, Dhillon, and Reddy, 2006). Hybrids with resistance to<br />

sorghum midge have been widely deployed in Australia for controlling sorghum midge,<br />

S. sorghicola (Henzell et al., 1997).<br />

Genetic Basis of Resistance<br />

Information on mechanisms and inheritance of resistance to insects can be utilized in<br />

selecting parents with diverse mechanisms or with different genes for resistance, selection<br />

of appropriate breeding methodology (pedigree, backcross, or population improvement)<br />

depending on the number of genes involved and nature of gene action, and developing<br />

isolines and mapping populations for resistance to insects. Resistance to insects may be<br />

oligogenic, polygenic, or cytoplasmic.<br />

Oligogenic Resistance<br />

Oligogenic resistance is controlled by one (monogenic or vertical resistance) or a few major<br />

genes, and the gene effects are easy to detect. Such resistance can be transferred into the<br />

elite lines through pedigree or backcross breeding. Resistance to several insects in rice,<br />

sweet potato, corn, and sorghum is monogenic in nature (Painter, 1958; M.D. Pathak, 1970;<br />

Kogan, 1982; H.C. Sharma, 1993). The chances of evolution of insect biotypes capable of<br />

overcoming the monogenic resistance are quite high, and such resistance breaks down<br />

over a period of time. Some of the notable examples of vertical resistance are resistance in

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