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Transgenic Resistance to Insects: Interactions with Nontarget Organisms 349<br />

less exposed to the transgene product as it is likely that not all of the prey will be contaminated.<br />

Therefore, it is necessary to differentiate them from the host-specifi c parasitoids,<br />

which are more likely to be affected by the toxins if the insect host acquires the toxins from<br />

the plants.<br />

Synergistic/Neutral Interactions<br />

Bt Toxins<br />

Predator levels in Bt cotton fi elds have been found to be signifi cantly greater than those<br />

in conventional cotton fi elds where insecticides were used for controlling the bollworm,<br />

H. armigera (Wu, Peng, and Jia, 2003). As the predator population increases, the outbreak<br />

of cotton aphid, A. gossypii in mid-season is controlled effectively, while the mirids become<br />

a major pest in Bt cotton because of reduced number of insecticide applications against<br />

H. armigera. Campylomma diversicornis Reuter, a dominant predator on eggs and newly<br />

hatched larvae of H. armigera, plays an important role in controlling insect pests in transgenic<br />

Bt cotton fi elds (W.X. Liu et al., 2000).<br />

No major differences have been observed in the abundance of generalist predators in<br />

fi elds with transgenic and nontransgenic crops (Hoffmann et al., 1992; Flint et al., 1995;<br />

Luttrell et al., 1995; Sims, 1995; Orr and Landis, 1997; Wang and Xia, 1997). Cui and Xia<br />

(1999) did not observe a signifi cant increase in populations of predatory arthropods in the<br />

Bt cotton fi eld, except that of Propylea japonica (Thun.). There was no apparent effect of<br />

transgenic cotton on the relative abundance of predatory spiders, Clubiona sp. and Neoscona<br />

sp., coccinellid, C. sexmaculatus, and the chrysopid, C. carnea. (Sharma, Pampapathy, and<br />

Arora, 2007). In Bt transgenic maize, temporal abundance of nontarget arthropods (bug,<br />

O. insidiosus; the syrphid, S. corollae; the ladybird, C. septempunctata; and the lacewing,<br />

C. carnea) varied greatly over the season, but did not differ between Bt and non-Bt maize<br />

(Bourguet et al., 2002). None of the coccinellids (three taxa), carnivorous carabids (three<br />

taxa), and ants were affected by cry3A-transgenic potatoes resistant to the Colorado potato<br />

beetle, L. decemlineata. Orius insidiosus was more abundant in transgenic fi elds than in nontransgenic<br />

fi elds in one year, while spiders were abundant in another year. The predator<br />

activity is not affected by pure transgenic and mixed seed potato crop (Riddick, Dively,<br />

and Barbosa, 1998). However, signifi cant effects on predator numbers were observed<br />

because of decline in the numbers of the insect hosts.<br />

There are no adverse effects of Bt maize on pre-imaginal development or mortality of<br />

C. carnea when reared on R. padi that had fed on Bt maize (Lozzia et al., 1998). Since there<br />

is no Bt toxin in the phloem as established by Raps et al. (2001), the aphids fed to the predators<br />

may have no Bt toxin, and thus no adverse effects of the transgenic plants could be<br />

measured. Dutton et al. (2002) observed that C. carnea larvae were not affected when fed<br />

spider mites, Tetranychus urticae Koch, that had been reared on Bt maize, even though the<br />

spider mites were found to contain more Cry1Ab toxin as compared to the lepidopteran<br />

larvae. Third instars of C. carnea showed a signifi cant preference for S. littoralis fed nontransgenic<br />

maize. However, no preference was displayed when C. carnea had the choice<br />

between R. padi fed on transgenic or nontransgenic maize. This lack of preference for<br />

R. padi fed on either transgenic or nontransgenic maize has been ascribed to the absence<br />

of the Bt protein in the phloem. In prey combinations with S. littoralis and R. padi, all three<br />

larval stages of C. carnea showed a preference for R. padi regardless of whether they had<br />

fed on transgenic or nontransgenic maize (Meier and Hilbeck, 2001). Romeis, Dutton, and<br />

Bigler (2004) suggested that C. carnea larvae are not sensitive to Cry1Ab, and that earlier

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