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

Molecular Markers and Metabolic Pathways<br />

There is considerable scope for changing the products of secondary metabolites that are<br />

associated with resistance to insect pests through biotechnological approaches. Harnessing<br />

synteny may have maximum benefi t where entire metabolic pathways are dissected and<br />

studied in detail in model systems, thereby identifying the key genes for manipulating<br />

that trait, which can then be traced in the species of interest. Many secondary plant metabolites<br />

such as fl avonoids, alkaloids, and terpenoids have been implicated in host plant<br />

resistance to insect pests. Many compounds of the fl avonoid biosynthetic pathway (fl avanones,<br />

fl avones, fl avanols, and isofl avonoids) accumulate in response to insect damage<br />

(Ebel, 1986; Sharma and Norris, 1991; Heller and Forkman, 1993). Molecular breeding and<br />

genetic engineering can be used to change the metabolic pathways to increase the amounts<br />

of various fl avonoids conferring resistance to insect pests, for example, medicarpin and<br />

sativan in alfalfa, cajanol and stilbene in pigeonpea, and stilbene in chickpea (Heller and<br />

Forkman, 1993). Stilbenes have been expressed in transgenic tobacco plants, exhibiting<br />

various degrees of inhibition of fungal growth (Heller and Forkman, 1993). Maysin, a glycosyl<br />

fl avone in maize silk, is associated with resistance to corn earworm, H. zea (Waiss<br />

et al., 1979). Most of the phenotypic variation in maysin concentration in maize silk is<br />

accounted for by the p1 locus, the transcription activator of the portion of the fl avonoid<br />

pathway leading to maysin synthesis. Reduced function p1 allele results in decreased transcription<br />

of genes encoding enzymes of the p1-controlled portion of the pathway, and thus<br />

reduced maysin synthesis. The marker umc105a corresponds to the brown pericarp (bp1)<br />

locus. The p1 and chromosome 9S regions are the major QTLs controlling silk antibiosis to<br />

the corm earworm (Byrne et al., 1997). Composite interval mapping has shown a major<br />

QTL in the asg20-whp1 interval of chromosome 2, and another near the wx1 locus on chromosome<br />

9 (Byrne et al., 1998). A gene that encodes chalcone synthase (whp1) on chromosome<br />

2 and a silk specifi c gene (sm1) on chromosome 6 affect silk maysin concentration<br />

and resistance to corn earworm in maize (Byrne et al., 1998). The extra chromosome 5A of<br />

Allium cepa L. plays an important role in fl avonoid biosynthesis (Masuzaki, Shigyo, and<br />

Yamauchi, 2006). The fl avonoid 3′-hydroxylase (F3′H) gene controlling quercetin synthesis<br />

from kaempferol is located on chromosome 7A, and an anonymous gene involved in<br />

glucosidation of quercetin is located on chromosome 3A or 4A.<br />

The Transgenic Approach and Gene Pyramiding through MAS<br />

Genetic engineering offers the advantage of rapid introgression of novel genes and traits<br />

into elite agronomic backgrounds (Mohan et al., 1997). Transgenic resistance to insects<br />

has been demonstrated in plants expressing insecticidal genes such as δ-endotoxins<br />

from Bacillus thuringiensis (Bt) Berliner, protease inhibitors, enzymes, secondary plant<br />

metabolites, and plant lectins (Sharma, Sharma, and Crouch, 2004). While transgenic<br />

plants with introduced Bt genes have been deployed in several crops on a global scale,<br />

the alternative genes have received considerably less attention. The potential of some of<br />

the alternative genes can only be realized by deploying them in combination with conventional<br />

host plant resistance and Bt genes (Sharma et al., 2002). Many of the candidate<br />

genes used in genetic transformation of crops are highly specifi c or are only mildly effective

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