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Mechanisms and Inheritance of Resistance to Insect Pests 129<br />

feeding by Spodoptera litura (F.), but result in slow growth of the larvae because of poor<br />

nutritional quality of the food or the presence of secondary plant substances (Stevenson<br />

et al., 1993). Larval mortality is the principal component of resistance to H. armigera in wild<br />

tomato, Lycopersicon hirsutum f. sp. glabratum Mull. (Tingey and Sinden, 1982; Kashyap and<br />

Verma, 1987). Antibiosis is also an important component of resistance to European corn<br />

borer, Ostrinia nubilalis (Hubner) and corn earworm, H. zea in maize (Klun, Tipton, and<br />

Brindley, 1967), and Colorado potato beetle, Leptinotarsa decemlineata (Say) (Sinden et al.,<br />

1986) and potato tuber moth, Phthorimaea operculella (Zeller) in potato. The larvae of cabbage<br />

looper, Trichoplusia ni (Hubner) are more effi cient in food utilization on the soybean<br />

susceptible genotype, Davis, than on the resistant genotype, PI 227687 (H.C. Sharma and<br />

Norris, 1993).<br />

Antibiosis expressed in terms of larval mortality, slow growth, and delayed development<br />

is one of the components of resistance to spotted stem borer, Chilo partellus (Swinhoe) (V.K.<br />

Kumar, Sharma, and Reddy, 2005), and the sorghum midge, S. sorghicola (H.C. Sharma,<br />

Vidyasagar, and Subramanian, 1993; H.C. Sharma, Franzmann, and Henzell, 2002) in sorghum.<br />

Antibiosis is also expressed in terms of reduced fecundity in the sorghum head bug,<br />

Calocoris angustatus (Lethiery) (H.C. Sharma, Lopez, and Nwanze, 1993).<br />

Tolerance<br />

Tolerance or recovery resistance enables a plant to withstand damage from an insect population<br />

that is injurious to another cultivar without a tolerance mechanism of resistance.<br />

Expression of tolerance is determined by the inherent capability of a genotype to overcome<br />

an insect infestation or to recover from insect damage and/or add new plant growth after<br />

insect damage. Plants with an ability to tolerate insect damage at times may produce more<br />

yield than the plants of a nontolerant susceptible cultivar at the same level of insect infestation.<br />

Tolerance often occurs in combination with antixenosis and antibiosis components<br />

of resistance. Production of tillers in sorghum following damage to the main plant by sorghum<br />

shoot fl y, A. soccata; and stem borer, C. partellu (H.C. Sharma and Nwanze, 1997),<br />

serves as a component of recovery resistance (Figure 5.5). Fertility status and moisture<br />

availability in the soil infl uence tiller production in plants damaged by shoot fl y and<br />

stem borers in sorghum (H.C. Sharma, 1993). Increase in grain mass in panicles partially<br />

damaged by sorghum midge, S. sorghicola (H.C. Sharma, 1997; H.C. Sharma, Abraham,<br />

and Stenhouse, 2002), and less grain damage per unit number of head bugs, C. angustatus<br />

(H.C. Sharma and Lopez, 1990, 1993; Padma Kumari, Sharma, and Reddy, 2000) serve as a<br />

component of recovery resistance to insect feeding and damage in sorghum. Temperature<br />

(Schweissing and Wilde, 1978) and nutrients (Schweissing and Wilde, 1979) affect the<br />

tolerance of sorghum seedlings to damage by greenbug, Schizaphis graminum (Rondani).<br />

Tolerance to insect damage has also been observed in alfalfa to weevil, Hypera postica<br />

(Gyllen.) (Dogger and Hanson, 1963), in maize to corn earworm, H. zea (Wiseman,<br />

McMillian, and Widstrom, 1972), in rice to brown planthopper, Nilaparvata lugens (Stal)<br />

(Panda and Heinrichs, 1983; Heinrichs et al., 1984), in wheat to greenbug, S. graminum<br />

(Starks and Merkle, 1977), in muskmelon to the aphid, Aphis gossypii Glover (Kennedy,<br />

Kishaba, and Bohn, 1975), and in turnip to cabbage maggot, Hylemyia sp. (Varis, 1958).<br />

Tolerance is an important component of resistance to H. armigera in cotton<br />

(Balasubramanian, Gopalan, and Subramanian, 1977; Murthy et al., 1998). It is expressed in<br />

terms of rejuvenation potential, healthy leaf growth, fl owering compensation potential,<br />

and plant vigor. The cotton genotype JK 276-4, possessing quick rejuvenation and higher<br />

fruiting effi ciency, suffers relatively less yield loss by H. armigera (Murthy et al., 1998).

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