Two–toxin strategies for management of insecticidal transgenic crops: can pyramiding succeed where pesticide mixtures have not?

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Abstract
Transgenic insect–resistant crops that express toxins from Bacillus thuringiensis (Bt) offer significant advantages to pest management, but are at risk of losing these advantages to the evolution of resistance in the targeted insect pests. All commercially available cultivars of these crops carry only a single Bt gene, and are particularly at risk where the targeted insect pests are not highly sensitive to the Bt toxin used. Under such circumstances, the most prudent method of avoiding resistance is to ensure that a large proportion of the pest population develops on non–transgenic ‘refuge’ hosts, generally of the crop itself. This has generated recommendations that 20% or more of the cotton and maize in any given area should be non–transgenic. This may be costly in terms of yields and may encourage further reliance on and resistance to pesticides. The use of two or more toxins in the same variety (pyramiding) can reduce the amount of refuge required to delay resistance for an extended period. Cross–resistance among the toxins appears to have been overestimated as a potential risk to the use of pyramids (and pesticide mixtures) because cross–resistance is at least as important when toxicants are used independently. Far more critical is that there should be nearly 100% mortality of susceptible insects on the transgenic crops. The past failures of pesticide mixtures to manage resistance provide important lessons for the most efficacious deployment of multiple toxins in transgenic crops.