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Development of Resistance to Transgenic Plants 391<br />

moth, P. xylostella as a model system to examine resistance management strategies. The<br />

higher number of larvae on refuge plants in fi eld tests indicated that a “separate refuge” is<br />

more effective in conserving susceptible larvae than a “mixed refuge,” and would thereby<br />

reduce the number of homozygous resistant (RR) offspring. Care must be exercised to<br />

ensure that refuges, particularly those sprayed with effi cacious insecticides, produce<br />

adequate numbers of susceptible insects.<br />

A strip confi guration is the least costly method of planting a 20% non-Bt maize refuge<br />

(Hyde et al., 2001). The non-Bt plants produce the susceptible insects, which have a probability<br />

of mating with insects emerging from the Bt crops nearby, and thus dilute the frequency<br />

of the resistant individuals. Separate refuges are superior to seed mixtures for<br />

delaying resistance. Movement of H. zea larvae from nontransgenic to transgenic plants<br />

may result in an increase in damage and reduce the yield in mixed stands of Bt and non-Bt<br />

plants (DuRant et al., 1996). In cotton, the numbers of eggs do not differ between mixed<br />

and pure stands of transgenic or nontransgenic plants (Halcomb et al., 1996; Lambert,<br />

Bradley, and Duyn, 1996). Planting two-row strips may be as good as separate refuges in<br />

delaying resistance, but their adoption carries greater risk because of the uncertainty surrounding<br />

the movement and survival of neonates (Onstad and Gould, 1998). Transgenic<br />

and nontransgenic plants could be grown in separate rows with a wider row spacing (strip<br />

planting) to minimize the rate of resistance development (Ramachandran et al., 1998a). As<br />

the proportion of nontransgenic plants increases, the number of larvae and amount of<br />

injury also increase.<br />

Role of Alternate Hosts as Refugia<br />

The ability of noncrops to support complete development of insect pests is an important<br />

factor in determining their impact on resistance management in transgenic crops. For<br />

polyphagous pests that feed on several fi eld crops and alternate hosts, there may not be<br />

any need to maintain refugia under subsistence farming conditions in the tropics, where<br />

several collateral and alternate hosts are available over time. There may not be any need to<br />

plant refuge crops for cereal stem borers and Heliothis/Helicoverpa in the tropics where they<br />

feed on several crop and weed hosts. The probability of European corn borer, O. nubilalis<br />

to complete development on a plant other than corn is relatively low, and the smallerstemmed<br />

noncorn plants may not make a substantial contribution to the pool of susceptible<br />

adults (Losey, Carter, and Silverman, 2002), although some individuals that survive could<br />

serve as a source of refuge. Cameron et al. (1997) studied the host range and occupancy of<br />

diamondback moth, P. xylostella, and potato tuber moth, P. operculella to determine what<br />

proportion of their populations occur in refugia and would therefore not be exposed to<br />

Bt-transformed Brassica or potato. Although P. operculella was found on Solanaceae other<br />

than potato in the fi eld, the alternative hosts were rare in areas where vegetables were<br />

growing and, therefore, were not a signifi cant source of refugia for this pest. Diamondback<br />

moth, P. xylostella, was common on Brassicaceae other than vegetable brassicas, particularly<br />

on white mustard, Sinapis alba L., which was the main refuge. Wild radish, Raphanus raphanistrum<br />

L., was less preferred, but was the most common alternative host plant. The area<br />

covered by wild radish was approximately 6% of the area planted in commercial brassicas<br />

and therefore formed a minor refuge for P. xylostella. The results suggested that to limit the<br />

development of resistance, insect-resistant transgenic potatoes should be less widely used<br />

than insect-resistant transgenic brassicas.<br />

A low proportion of potato tuber moths, P. operculella, foraged beyond 100 to 250 m to<br />

infest tubers or plants (Cameron et al., 2002). Light traps indicated that the number of

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