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134 Biotechnological Approaches for Pest Management and Ecological Sustainability<br />

wheat to Hessian fl y, M. destructor (Smith, 2005), and rice resistance to brown planthopper,<br />

N. lugens (Khush and Brar, 1991). The vertical type of resistance for tolerance or moderate<br />

levels of resistance provides stable resistance to insect pests (Panda and Khush, 1995). Since<br />

availability of new genes for resistance may be a limiting factor, it is important to manage<br />

known resistant genes to maximize their effect and durability through: (1) recycling and<br />

sequential release of resistance genes, (2) gene pyramiding, (3) regional biotype-specifi c<br />

deployment of resistance genes, and (4) development of synthetics involving multilines<br />

with different genes for resistance.<br />

Polygenic Resistance<br />

Polygenic resistance is controlled by several genes. It is also called horizontal resistance.<br />

Inheritance of such genes is usually complex, and it takes a long time to transfer polygenic<br />

resistance into improved cultivars. Pedigree breeding involving multiple crosses or population<br />

improvement approaches can be used to breed for horizontal resistance. Because<br />

of the additive effect of several genes, the insects take a longer time to overcome such<br />

resistance and thus the chances of evolution of new insect biotypes are low or minimal.<br />

Resistance to several insect species in crops such as cotton, legumes, maize, rice, and wheat<br />

is under polygenic control (Panda and Khush, 1995; Smith, 2005). Progress in breeding for<br />

insect resistance involving a horizontal or additive type of gene action has been slow, and<br />

suffi cient levels of resistance have not been achieved in cultivars with desirable agronomic<br />

backgrounds. The horizontal type of resistance does not exert a selection pressure on pest<br />

populations, and is quite durable. However, strong environmental infl uence and genetic<br />

variability in insect populations complicate the identifi cation and transfer of resistance<br />

genes into high-yielding cultivars.<br />

Selection for the additive type of gene action should be deferred until later generations,<br />

when a desired level of homozygosity is achieved. In the early generations, selection may be<br />

exercised against lines highly susceptible to the target insect pests. Indirect selection using<br />

physico-chemical traits associated with insect resistance can also be practiced in the early<br />

generations, for example, glossy leaf trait for resistance to shoot fl y, A. soccata and short and<br />

tight glumes for resistance to midge, S. sorghicola in sorghum (H.C. Sharma and Nwanze,<br />

1997), trichomes for H. zea resistance in tomato (Farrar and Kennedy, 1987), leaf hairs for<br />

resistance to leafhopper, Amrasca biguttula biguttula Ishida in cotton (H.C. Sharma and Agarwal,<br />

1983), and nonglandular trichomes in wild relatives of pigeonpea, Cajanus scarabaeoides (L.)<br />

F. Muell. for resistance to H. armigera (Romeis et al., 1999; Rupakala et al., 2005). Molecular<br />

markers linked with insect resistance genes or the factors associated with insect resistance<br />

can be used to improve the effi ciency of screening and breeding for resistance to insects,<br />

including the additive type of gene action (H.C. Sharma, Abraham, and Stenhouse, 2002).<br />

Cytoplasmic Effects<br />

Cytoplasmic resistance is due to the factors in the cytoplasm of the host plant. Cytoplasmic<br />

inheritance is maternal, and can be detected by making reciprocal crosses. The fi rst cytoplasmic<br />

male-sterile line in rice was developed by substituting nuclear genes of the indica<br />

variety, Taichung Native 1 (Athwal and Virmani, 1972), and since then, a large number of<br />

cytoplasmic male-sterile (CMS) lines have been developed by exploiting both intra- and<br />

interspecifi c CMS systems (Virmani and Wan, 1988). The male sterile cytoplasm affects rice<br />

plant reactions to pathogens, as the WA male sterile cytoplasm in rice is less susceptible to<br />

Pyricularia oryzae Cav. and Xanthomonas oryzae pv. oryzae (Swings et al.) than the fertile<br />

cytoplasm (R.C. Yang et al., 1989). In maize, the use of CMS-T cytoplasm for hybrid production

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