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

Acetone extracts of pods of pigeonpea and its wild relative, Cajanus platycarpus (Benth.)<br />

Maesen, have a signifi cant feeding stimulant effect on H. armigera larvae, whereas the<br />

extracts from C. scarabaeoides pods have no such effect (Shanower, Yoshida, and Peter, 1997).<br />

Quercetin, quercetrin, and guercetin-3-methyl ether (present in pod surface exudates) play<br />

an important role in food selection behavior of H. armigera larvae in pigeonpea (Green<br />

et al., 2002, 2003). Sterols and soybean leaf extractables in combination with sucrose are<br />

phagostimulant to the larvae of the cabbage looper, T. ni (Sharma and Norris, 1994a).<br />

Antifeedants<br />

Chemicals that inhibit feeding are called phagodeterrents. High acidity in the leaf exudates<br />

of chickpea is associated with resistance to H. armigera (Srivastava and Srivastava,<br />

1989; Bhagwat et al., 1995). Chickpea exudates have malate and oxalate as the main components,<br />

and there are characteristic differences among genotypes depending on diurnal<br />

cycles and growth stage. Varieties with the highest amounts of malic acid are resistant to<br />

H. armigera (Rembold, 1981; Rembold et al., 1990). Malic acid acts as an antifeedant to the<br />

H. armigera larvae (Bhagwat et al., 1995). Oxalic acid inhibits the growth of H. armigera<br />

larvae when incorporated into artifi cial diet, while malic acid shows no growth inhibition<br />

(Yoshida, Cowgill, and Wightman, 1995, 1997). The chickpea fl avonoids judaicin 7-O- -<br />

glucoside, 2 methoxy judaicin, judaicin, and maakiain present in wild relatives of chickpea<br />

(Cicer bijugum Rech. and C. judaicum Boiss.) have shown antifeedant activity towards the<br />

larvae of H. armigera (Simmonds and Stevenson, 2001). Stilbene, a phytoalexin, occurs at high<br />

concentrations in pigeonpea cultivars with resistance to H. armigera (Green et al., 2003).<br />

Growth Inhibitors<br />

Alkaloids, ketones, terpenoids, fl avonoids, and organic acids produced by the plants are<br />

toxic to insects. Gossypol, heliocide 1, heliocide 2, gossypolone, hemigossypolone, lactone,<br />

condensed tannins, and volatile terpenes in cotton confer resistance to bollworms (Hedin<br />

et al., 1983; Sharma, 1982; Sharma and Agarwal, 1982a, 1982b; Tang and Wang, 1996). High<br />

density of gossypol glands on squares rather than leaves, calyxes, and bracts contributes to<br />

H. armigera resistance in cotton (Rajarajeswari and Subba Rao, 1997). Tannins and terpenoid<br />

aldehydes gradually increase from the cotyledon stage to the late-bloom stage of cotton<br />

(Zummo, Segers, and Benedict, 1984). However, there is a sudden drop in tannins at the<br />

one-third growth stage, which provides a window of increased vulnerability to H. zea.<br />

Phenolics have also been reported to impart resistance to bollworms. Catechin, chrysanthemin,<br />

isoquercitrin, quercetin, condensed tannins, cyanidin, and delphinidin have been<br />

tested in laboratory bioassays against H. armigera, and found to exert a weight inhibition<br />

of 50% at variable concentrations. The polar solvent extractables of the soybean genotype<br />

PI 227687, resistant to the cabbage looper, T. ni, contains diadzien, coumesterol, sojagol, and<br />

glyceollins. These compounds reduce feeding, survival, and development of T. ni (Sharma<br />

and Norris, 1991, 1994b).<br />

In tomato, L-tomatine, 2-tridecanone, and phenolics are responsible for antibiosis to<br />

H. zea (Ferry and Cuthbert, 1975; Dimock and Kennedy, 1983; Farrar and Kennedy, 1988,<br />

1990). The allelochemical 2- tridecanone is highly toxic to H. zea (Kennedy and Yamamoto,<br />

1979). High phenol content has also been reported to be associated with resistance to<br />

H. armigera (Banerjee and Kalloo, 1989). Acylglucoses exuded by trichomes of the wild<br />

tomato species, Lycopersicon pennellii (Corr.) D’Arcy act as feeding or oviposition deterrents<br />

or both. Exposure to these compounds in artifi cial diet or as sprays on cultivated tomato

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