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

dynamics and movement of insects between transgenic and refuge fi elds on evolution of<br />

insecticide resistance have been examined in two different simulation models (Caprio,<br />

2001). The two models were developed to test the hypothesis that increasing the habitat<br />

from fi ne-grained to coarse-grained, and the resultant increase in nonrandom mating,<br />

would increase the rate of local adaptation and hence evolution of resistance. The fi rst, a<br />

stochastic, spatially explicit model, altered habitat by varying adult dispersal rates between<br />

habitat patches. In contrast to the expectation that increasing patch isolation and increasing<br />

the coarseness of the habitats would increase the rate of resistance evolution, intermediate<br />

levels of dispersal actually delayed resistance by as much as fi ve-fold over the range<br />

of dispersal levels observed. A simple deterministic model was also developed to separate<br />

the impact of mating and ovipositional behavior. This model showed qualitatively the<br />

same results. Under similar assumptions, it predicted longer delays in resistance evolution.<br />

In this model, nonrandom mating alone increased the rate at which insects adapted<br />

to transgenic crops, but nonrandom mating in combination with nonrandom oviposition<br />

could signifi cantly delay resistance evolution. Differences between the two models may be<br />

due to population regulation incorporated in a spatially explicit model. The models clearly<br />

suggested that resistance management programs using untreated refuges should not overemphasize<br />

random mating at the cost of making the habitat too fi ne-grained.<br />

Since resistance gene(s) do not appear to be completely recessive, the need for a higher<br />

proportion of refuge vis-à-vis transgenic crop has often been advocated (Gould and<br />

Tabashnik, 1998). The resistant individuals emerging out of Bt cotton should mate randomly<br />

with those from the refuge where the susceptible individuals breed. However, synchrony<br />

of emergence of adults from Bt cotton and refuge will be equally essential. Helicoverpa<br />

armigera, a highly mobile insect, may mate randomly, although closeness of refuge as a<br />

policy also contributes to its success. Soybean and peanut as mixed crops could provide<br />

natural refuge for the second to fourth generations of H. armigera in North China (Wu,<br />

Guo, and Gao, 2002). However, function of refuge appeared to depend on the proportion of<br />

Bt cotton. However, there is a need to protect the natural refuge of noncotton host plants of<br />

H. armigera by not commercializing both Bt cotton and Bt corn in the same area (Zhao<br />

et al., 2000b). In India, small farm holdings may not necessarily follow refuge norms. As<br />

the transgenic cotton area increases, it may lead to almost monocropping without a signifi -<br />

cant contribution of mixed crops or nontrangenic Bt cotton towards the production of<br />

susceptible individuals.<br />

Conclusions<br />

The use of crop protection traits through transgenics will continue to expand in the future,<br />

and gene pyramiding might become very common. This approach of controlling insects<br />

would offer the advantage of allowing some degree of selection for specifi city effects,<br />

so that pests but not benefi cial organisms are targeted. Resistance to Bt is a recessive or<br />

partially recessive mutation in a major autosomal gene, and cross resistance extends to Cry<br />

proteins sharing the same binding site(s). Cry proteins with specifi city to different binding<br />

sites should be considered for gene pyramiding as a component of a resistance management<br />

strategy. In a vast majority of cases, resistance has been found to be quite unstable<br />

because of fi tness costs associated with resistance, although there are a few exceptions. A<br />

detailed understanding of the mechanisms, insect biology, and plant molecular biology

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