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

only 2.3-fold resistance to Cry2Aa, suggesting that Cry1Ac resistance extended to Cry1Ab,<br />

but not to Cry2Aa. Helicoverpa armigera resistant to Cry1Ac did not possess cross resistance<br />

to Cry1C and Cry2Aa, but was resistant to Bt corn expressing Cry1A (Zhao et al., 2000b).<br />

Chandrashekar et al. (2005) also observed 31.4-fold resistance to Cry1Ac, 25.4-fold resistance<br />

to Cry1Ab, and only 8.4-fold resistance to Cry1Aa in strains of H. armigera selected<br />

for resistance to these Bt toxins.<br />

Resistance to the Bt formulation, Javelin increased 1.9 to 4.4 times under fi eld conditions,<br />

but was signifi cant only at an application rate of 1.12 kg ha 1 , irrespective of the presence or<br />

absence of a refuge (Perez, Shelton, and Roush, 1997). Selection with Javelin at 0.3 kg ha 1<br />

or Xentari did not cause a signifi cant increase in resistance to Btk nor did P. xylostella<br />

selected with Xentari resistance show resistance to B. thuringiensis subsp. aizawai. Two strains<br />

of the diamondback moth, P. xylostella, selected using Cry1C protoxin and transgenic broccoli<br />

plants expressing cry1C, were resistant to Cry1C, but had different cross-resistance<br />

patterns (Zhao et al., 2001). In a study involving 12 protoxins (Cry1Aa, Cry1Ab, Cry1Ac,<br />

Cry1Bb, Cry1C, Cry1D, Cry1E, Cry1F, Cry1J, Cry2Ab, Cry9Aa, and Cry9C), the resistance<br />

ratio (RR) of one strain (BCS-Cry1C-1) to the Cry1C protoxin was 1,090-fold with a high<br />

level of cross-resistance to Cry1Aa, Cry1Ab, Cry1Ac, Cry1F, and Cry1J (RR 390-fold). The<br />

cross resistance to Cry1A, Cry1F, and Cry1J in this strain was probably related to Cry1A<br />

resistance gene(s) that came from the initial fi eld population as a result of intensive sprayings<br />

of Bt products containing Cry1A protoxins. The neonates of this strain survived on<br />

transgenic broccoli plants expressing either cry1Ac or cry1C toxins. The other strain<br />

(BCS-Cry1C-2) resistant to Cry1C did not show cross resistance to other Bt protoxins. The<br />

neonates of this strain survived on transgenic broccoli expressing cry1C, but not cry1Ac.<br />

Genetics of Resistance<br />

Frequency of Resistance Alleles<br />

Frequency of resistance allele between generations in the insect population is based on the<br />

selection intensity, fi tness cost, percentage of refugia, mortality in refugia due to insecticide<br />

sprays, initial frequency of the resistance allele, and the rate of insect movement<br />

(Cerda and Wright, 2004). A simulation model for resistance development based on four<br />

different spatial patterns of refugia inside the fi eld (border, central, equidistant, and random),<br />

four crop rotation patterns, fi ve different sizes of refuge (5% to 50%), and two levels<br />

of non-Bt insecticide plus an untreated control in the refugia indicted that:<br />

• The greater the size of refugia, the lower the rate of increase in the frequency of<br />

resistance alleles.<br />

• Positioning patches of refugia at random in the fi eld resulted in a higher rate of<br />

increase in the frequency of resistance alleles compared with nonrandom<br />

positions.<br />

• Positioning refugia along the border resulted in a lower rate of increase in the<br />

frequency of resistance alleles compared with equidistant and central patterns.<br />

• The frequency of resistance allele increased markedly when the refugia was less<br />

than 10%.

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