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Physico-Chemical and Molecular Markers for Resistance to Insect Pests 165<br />

leaves resulted in retarded growth rates, extended development period, and reduced survival<br />

of H. zea and S. exigua ( Juvik et al., 1994). In the case of H. virescens resistance in<br />

tobacco, larval weight gain is negatively correlated with nicotine content at various leaf<br />

positions. Larvae survived better and grew faster on the bud leaves, which have lower<br />

nicotine levels than the leaves further down the stalk where alkaloid levels are high<br />

( Jackson, Johnson, and Stephenson, 2002). Luteolin-C-glycosides, maysin, and isomaysin,<br />

which occur in maize silks, confer resistance to H. zea (Waiss et al., 1979). In soybean, pinitol<br />

confers resistance to H. zea (Dougherty, 1976), while long chain fatty acids in sunfl ower<br />

act as growth inhibitors for H. virescens and H. zea (Elliger et al., 1976). In potato, concentrations<br />

of the glycoalkaloids, solanine and chaconine are associated with resistance to<br />

E. fabae (Sanford et al., 1992).<br />

Nonprotein Amino Acids<br />

Nonprotein or unusual amino acids are found in a number of plant species, and afford<br />

protection against herbivores. The protective effect is elicited through their structural analogy<br />

to the commonly occurring essential amino acids. The harmful effects on insects occur<br />

as these get incorporated into proteins that are toxic to insects. Among these, L-canavanine,<br />

azetidine-2-caboxylic acid, 2,4-diamino butyric acid, minosine, and 3-hydroxyproline have<br />

signifi cant growth inhibition effects on insects (Parmar and Walia, 2001). L-canavanine is<br />

a structural homologue of L-arginine, and occurs in over 1,500 leguminous plant species.<br />

Some of the nonprotein amino acids also act as enzyme inhibitors. Canaline, a hydrolytic<br />

product of canavanine, inhibits pyridoxal phosphate-dependent enzymes by forming a<br />

covalent bond. Minosine is a strong inhibitor of rat liver cytothionine synthetase and cytothionase,<br />

and inhibits the growth of rust red fl our beetle, Tribolium castaneum (Herbst)<br />

(Ishaaya et al., 1991).<br />

Nutritional Factors<br />

For a plant to serve as a host for an insect, it should be able to provide holistic nutrition to<br />

support growth, development, and reproduction (Beck, 1972). It should also be suitable for<br />

digestion, assimilation, and conversion into energy and structural compounds required by<br />

the insects. Some of the nutrients such as amino acids, phospholipids, fatty acids, steroids,<br />

and ascorbic acid also serve as phagostimulants for different insect species (Dadd, 1970).<br />

Variation in the amounts of these compounds results in various degrees of host plant<br />

acceptance and damage by the herbivorous insects. Imbalanced ascorbic acid content in<br />

maize (Penny, Scott, and Guthrie, 1967), amino acid content in pea (Auclair, Maltais, and<br />

Cartier, 1957), low quantities of glutamic acid and asparagine in rice (Sogawa and Pathak,<br />

1970), low lysine content in sorghum (Singh and Jotwani, 1980), and high proline content in<br />

cotton (Sharma and Agarwal, 1983a) are associated with resistance to insects. Total soluble<br />

sugars in the pod wall have a signifi cant and negative correlation with pod damage by<br />

H. armigera in pigeonpea. Pea varieties defi cient in certain amino acids are resistant to the<br />

pea aphid, Acyrthosiphon pisum (Harris) (Auclair, 1963).<br />

In cotton, higher amounts of reducing sugars are associated with resistance to the<br />

leafhopper, A. biguttula biguttula (Singh and Agarwal, 1988), and amino acid content to<br />

stem weevil, Pempherulus affi nis (Kalt.) (Parameswaran, 1983). Rice varieties resistant to<br />

thrips, Stenchaetothrips biformis (Bagnall), have lower amounts of sugars and free amino<br />

acids (Thayumananan et al., 1990), while low amounts of amino acids are associated with<br />

resistance to brown planthopper, N. lugens (Sogawa and Pathak, 1970; Bharathi, 1989).

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