Evolutionary ecology of insect adaptation to Bt crops

Abstract
Transgenic crops producing Bacillus thuringiensis (Bt) toxins are used worldwide to control major pests of corn and cotton. Development of strategies to delay the evolution of pest resistance to Bt crops requires an understanding of factors affecting responses to natural selection, which include variation in survival on Bt crops, heritability of resistance, and fitness advantages associated with resistance mutations. The two main strategies adopted for delaying resistance are the refuge and pyramid strategies. Both can reduce heritability of resistance, but pyramids can also delay resistance by reducing genetic variation for resistance. Seasonal declines in the concentration of Bt toxins in transgenic cultivars, however, can increase the heritability of resistance. The fitness advantages associated with resistance mutations can be reduced by agronomic practices, including increasing refuge size, manipulating refuges to increase fitness costs, and manipulating Bt cultivars to reduce fitness of resistant individuals. Manipulating costs and fitness of resistant individuals on transgenic insecticidal crops may be especially important for thwarting evolution of resistance in haplodiploid and parthenogenetic pests. Field-evolved resistance to Bt crops in only five pests during the last 14 years suggests that the refuge strategy has successfully delayed resistance, but the accumulation of resistant pests could accelerate.