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

Schwarzbach, and Fischbeck, 1983). Recent reports on the duration of induced resistance in<br />

host-pathogen systems have been documented in barley (Pelcz and Wolffgang, 1986), cotton<br />

(Liu, Chen, and Wang, 1990), and cucumber (Dalisay and Kuc, 1995). Insect herbivory<br />

has been shown to induce resistance in soybean plants (Lin, Kogan, and Fischer, 1990;<br />

Fischer, Kogan, and Paxton, 1990; Lin and Kogan, 1990) against the soybean looper,<br />

Pseudoplusia includens (Walker) and the Mexican bean beetle, E. varivestis. Cross-resistance<br />

to P. includens due to previous Cerotoma trifurcata (Forster) injury in the form of induced<br />

resistance is benefi cial to the plant. Bean leaf beetle, C. trifurcata herbivory also affects<br />

larval growth rates and reduced the suitability of foliage to the corn earworm, H. zea in<br />

soybean (Felton, Summers, and Mueller, 1994).<br />

Chemically induced expression systems or “gene switches” enable temporal, spatial, and<br />

quantitative control of genes introduced into plants or those that are already present in the<br />

plants to impart resistance to insects. This approach has provided opportunities for management<br />

of development of resistance in insect pest populations in transgenic crops. In addition<br />

to insect or pathogen attack, resistance can also be induced by suboptimal concentrations of<br />

potassium iodide, copper, and herbicides. Effectiveness of the chemical injury inducer<br />

Actigard in providing resistance to various insect pests and pathogens in the tomato has<br />

been demonstrated by Inbar et al. (1998). Induced resistance against E. varivestis lasted three<br />

days after damage in soybean (Underwood, 1998). Proteinase inhibitors and oxidative<br />

enzymes such as polyphenol oxidase, peroxidase, and lipoxygenase persist for at least<br />

21 days after induction in damaged tomato leafl ets (Stout, Workman, and Duffey, 1996).<br />

A wide range of inducible genes has been identifi ed in plants based on endogenous<br />

chemical signals, such as phytohormones, responses to insect and pathogen attack, or<br />

wounding. The best-studied system utilizes the PR1-a promoter from tobacco, which is<br />

induced during systemic resistance response following pathogen infection (Uknes et al.,<br />

1993). The PR1-a mRNA levels can also be induced by exogenous application of salicylic<br />

acid or 1,2,3-benzothaidiazole-7-carbothioic acid S-methyl ester (BTH) (Ward et al., 1991;<br />

Gatz, 1997). The latter has been commercialized as a plant immunization chemical.<br />

However, PR1-a use may be limited due to its responsiveness to exogenous signals such as<br />

pathogens, UV-B, and pollutants (Gatz, 1997). Safeners used for detoxifi cation of certain<br />

herbicides also induce a range of metabolic enzymes, such as glutathione-S transferase and<br />

cytochrome P 450 ( Jepson, Martinez, and Sweetruan, 1998). Safener-induced cDNA clones<br />

(In 2-2 and GST-27) have been isolated from maize and Arabidopsis. These promoters can be<br />

induced by M-(aminocarbonyl)-2-chloro benzenesulfonamide and the herbicide sulfonylurea.<br />

Tetracycline and lactose represser operator systems from Escherichia coli Escherich have<br />

also been used to control gene expression ( Jepson et al., 1994). Tetracycline (TetR) repressor<br />

protein has been used to regulate the expression from CaMV 35S promotor (Gatz and<br />

Quail, 1998). Another system in yeast uses a copper-dependent transcriptional activation<br />

system, which consists of ace1 gene encoding a metalloresponsive factor expressed constitutively.<br />

Activation of the reporter gene (ace1) is achieved in the presence of copper (Mett,<br />

Lochhead, and Reynolds, 1993). Copper has traditionally been used as a fungicide, but<br />

application of copper for activation of the reporter gene may not be ideal in all crops.<br />

Another transcriptional activation system (Alc) is based on ethanol (Caddick et al., 1998).<br />

The AlcR transcription factor interacts with ethanol or related inducers. It has a low activity,<br />

rapid and reversible induction, and suffi ciently high levels of induction to generate<br />

phenotypic effects. Glucocorticoid receptor (GR) (Wang et al., 1997) and ecdysone receptor<br />

(Wing, Slawecki, and Carlson, 1988; Jepson, Martinez, and Sweetruan, 1998) also have the<br />

potential for use as switches for inducible resistance to insects.<br />

Octadecanoid and the salicylic acid pathways are involved in the induced attraction of the<br />

parasitoid wasp, Cotesia rubecula (Marshall) by Arabidopsis thaliana (L.) Heyn. infested with

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