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Transgenic Resistance to Insects: Interactions with Nontarget Organisms 341<br />

measure the ecological effects of transgenic plants on nontarget organisms. Endogenous<br />

production of Bt toxins in all parts of the plant throughout the crop-growing season may<br />

lead to a different mode of exposure of herbivores, and thus their parasites and predators.<br />

Also, altered metabolism of plants (by virtue of production of toxin protein or insertion of<br />

the novel gene) may also affect the interaction between the plants, insects, and their natural<br />

enemies. Therefore, there is a need for long-term studies involving insect-resistant<br />

transgenic plants, including lethal, sublethal, and chronic effects in nontarget organisms.<br />

Bt Sprays, Transgenic Plants, and Nontarget Organisms<br />

The Bt spores and toxins are degraded quickly. Under UV light, the toxins decay within a<br />

few hours to a few days, while the spores need soil for germination (Schutte and Riede,<br />

1998). In comparison, the toxin proteins in transgenic plants are produced throughout the<br />

crop cycle in all parts of the plant. Formulations based on Bt have shown toxic effects<br />

against some nontarget insects (Wagner et al., 1996), but only a few of the parasitoids and<br />

predators may be directly harmed by Bt sprays (Flexner, Lighthart, and Croft, 1986; Deml<br />

and Dettner, 1998). Application of Bt as foliar sprays does not cause any harmful effects to<br />

microorganisms or to vertebrates (Kreutzweiser et al., 1996). Bt sprays have shown negative<br />

effects against 24 species each of parasites and predators, but no adverse effects have<br />

been observed against 78 predators and 87 parasitoids (Deml and Dettner, 1998). Indirect<br />

effects through nutritional quality of the prey have been observed on 12 predators and<br />

eight parasitoids. High levels of Bt toxicity have been observed only against two species,<br />

but low levels of toxicity have been observed against 29 species. Toxins from B. thuringiensis<br />

var. tenebrionis have shown no effects against 9 of the 11 species examined, but high<br />

mortality and reduced fecundity have been observed in two species. Under fi eld conditions,<br />

Newleaf (Cry3Aa) potato plants have been found to be highly effective in suppressing<br />

populations of Colorado potato beetle, Leptinotarsa decemlineata (Say) and provided<br />

better control than weekly sprays of a microbial Bt-based formulation containing Cry3Aa,<br />

without reducing the number of the generalist predators, and thus is compatible with integrated<br />

pest management (IPM) programs (Reed et al., 2001).<br />

Transgenic plants express semiactivated proteins, which are ingested by the insects. In<br />

this process, no crystal solubilization and protoxin-toxin conversion is necessary in the<br />

insect midgut. Increased activity has been observed in some herbivorous insects when fed<br />

with toxin instead of the protoxin, suggesting that such insects lack the appropriate<br />

enzyme composition or pH that converts the protoxin to toxin (Moar et al., 1995). The truncated<br />

and activated toxins expressed in transgenic plants constitutively in high doses kill<br />

the insects much faster than the Bt sprays, which need to be activated by the insect enzymes.<br />

Arthropod species that have the binding sites but cannot activate the toxins could be<br />

harmed by the toxins from the transgenic plants. However, Bt toxins can be activated in<br />

different ways, leading to a change in their specifi city (Haider, Knowles, and Ellar, 1986).<br />

While Bt sprays might affect all the insects in an ecosystem wherever they are sprayed, the<br />

toxins from the transgenic plants affect only the insects that ingest the toxins from the<br />

plants through feeding or indirectly through the insect host or prey (Figure 11.1). Therefore,<br />

toxicity assays from Bt sprays cannot be substituted for the toxins expressed in transgenic<br />

plants. The Bt protein molecule is quite complex, and its mode of action on nontarget<br />

organisms is not understood properly. Little information is available on mode of action in

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