Effects of cotton cultivar on fitness costs associated with resistance of pink bollworm (Lepidoptera: Gelechiidae) to Bt cotton

Fitness costs associated with insect resistance to transgenic crops producing toxins from Bacillus thuringiensis (Bt) reduce the fitness on non-Bt refuge plants of resistant individuals relative to susceptible individuals. Because costs may vary among host plants, choosing refuge cultivars that increase the dominance or magnitude of costs could help to delay resistance. Specifically, cultivars with high concentrations of toxic phytochemicals could magnify costs. To test this hypothesis, we compared life history traits of three independent sets of pink bollworm, Pectinophora gossypiella (Saunders), populations on two cotton cultivars that differed in antibiosis against this cotton pest. Each set had an unselected susceptible population, a resistant population derived by selection from the susceptible population, and the F1 progeny of the susceptible and resistant populations. Confirming previous findings with pink bollworm feeding on cotton, costs primarily affected survival and were recessive on both cultivars. The magnitude of the survival cost did not differ between cultivars. Although the experimental results did not reveal differences between cultivars in the magnitude or dominance of costs, modeling results suggest that differences between cultivars in pink bollworm survival could affect resistance evolution. Thus, knowledge of the interaction between host plants and fitness costs associated with resistance to Bt crops could be helpful in guiding the choice of refuge cultivars.     

Fitness cost of resistance to Bt cotton linked with increased gossypol content in pink bollworm larvae

Fitness costs of resistance to Bacillus thuringiensis (Bt) crops occur in the absence of Bt toxins, when individuals with resistance alleles are less fit than individuals without resistance alleles. As costs of Bt resistance are common, refuges of non-Bt host plants can delay resistance not only by providing susceptible individuals to mate with resistant individuals, but also by selecting against resistance. Because costs typically vary across host plants, refuges with host plants that magnify costs or make them less recessive could enhance resistance management. Limited understanding of the physiological mechanisms causing fitness costs, however, hampers attempts to increase costs. In several major cotton pests including pink bollworm (Pectinophora gossypiella), resistance to Cry1Ac cotton is associated with mutations altering cadherin proteins that bind this toxin in susceptible larvae. Here we report that the concentration of gossypol, a cotton defensive chemical, was higher in pink bollworm larvae with cadherin resistance alleles than in larvae lacking such alleles. Adding gossypol to the larval diet decreased larval weight and survival, and increased the fitness cost affecting larval growth, but not survival. Across cadherin genotypes, the cost affecting larval growth increased as the gossypol concentration of larvae increased. These results suggest that increased accumulation of plant defensive chemicals may contribute to fitness costs associated with resistance to Bt toxins.     

Effects of four nematode species on fitness costs of pink bollworm resistance to Bacillus thuringiensis toxin Cry1Ac

Evolution of resistance by pests can reduce the efficacy oftransgenic crops that produce insecticidal toxins from the bacterium Bacillus thuringiensis Berliner (Bt). In conjunction with refuges of non-Bt host plants, fitness costs can delay the evolution of resistance. Furthermore, fitness costs often vary with ecological conditions, suggesting that agricultural landscapes can be manipulated to magnify fitness costs and thereby prolong the efficacy of Bt crops. In the current study, we tested the effects of four species of entomopathogenic nematodes (Steinernematidae and Heterorhabditidae) on the magnitude and dominance of fitness costs of resistance to Bt toxin CrylAc in pink bollworm, Pectinophora gossypiella (Saunders) (Lepidoptera: Gelechiidae). For more than a decade, field populations of pink bollworm in the United States have remained susceptible to Bt cotton Gossypium hirsutum L. producing CrylAc; however, we used laboratory strains that had a mixture of susceptible and resistant individuals. In laboratory experiments, dominant fitness costs were imposed by the nematode Steinernema riobrave Cabanillas, Poinar, and Raulston but no fitness costs were imposed by Steinernema carpocapsae Weiser, Steinernema sp. (ML18 strain), or Heterorhabditis sonorensis Stock, Rivera-Ordu?o, and Flores-Lara. In computer simulations, evolution of resistance to Cry1Ac by pink bollworm was substantially delayed by treating some non-Bt cotton refuge fields with nematodes that imposed a dominant fitness cost, similar to the cost observed in laboratory experiments with S. riobrave. Based on the results here and in related studies, we conclude that entomopathogenic nematodes could bolster insect resistance management, but the success of this approach will depend on selecting the appropriate species of nematode and environment, as fitness costs were magnified by only two of five species evaluated and also depended on environmental factors.     

Fitness costs associated with Cry1F resistance in the European corn borer

Crops producing insecticidal toxins derived from the bacterium Bacillus thuringiensis (Bt) are widely planted to manage insect pests. Bt crops can provide an effective tool for pest management; however, the evolution of Bt resistance can diminish this benefit. The European corn borer, Ostrinia nubilalis Hübner, is a significant pest of maize and is widely managed with Bt maize in the Midwest of the United States. When Bt crops are grown in conjunction with non‐Bt refuges, fitness costs of Bt resistance can delay the evolution of resistance. Importantly, fitness costs often vary with ecological factors, including host‐plant genotype and diapause. In this study, we examined fitness costs associated with Cry1F resistance in O. nubilalis when insects were reared on three maize lines. Fitness costs were tested in two experiments. One experiment assessed the fitness costs when Cry1F‐resistant and Cry1F‐susceptible insects were reared on plants as larvae and experienced diapause. The second experiment tested resistant, susceptible and F1 heterozygotes that were reared on plants but did not experience diapause. Despite some evidence of greater adult longevity for Cry1F‐resistant insects, these insects produced fewer fertile eggs than Cry1F‐susceptible insects, and this occurred independent of diapause. Reduced fecundity was not detected among heterozygous individuals, which indicated that this fitness cost was recessive. Additionally, maize lines did not affect the magnitude of this fitness cost. The lower fitness of Cry1F‐resistant O. nubilalis may contribute to the maintenance of Cry1F susceptibility in field populations more than a decade after Cry1F maize was commercialized.     

Insect resistance to transgenic Bt crops: lessons from the laboratory and field

Transgenic crops that produce insecticidal toxins from the bacterium Bacillus thuringiensis (Bt) grew on >62 million ha worldwide from 1996 to 2002. Despite expectations that pests would rapidly evolve resistance to such Bt crops, increases in the frequency of resistance caused by exposure to Bt crops in the field have not yet been documented. In laboratory and greenhouse tests, however, at least seven resistant laboratory strains of three pests (Plutella xylostella [L.], Pectinophora gossypiella [Saunders], and Helicoverpa armigera [Hübner]) have completed development on Bt crops. In contrast, several other laboratory strains with 70- to 10,100-fold resistance to Bt toxins in diet did not survive on Bt crops. Monitoring of field populations in regions with high adoption of Bt crops has not yet detected increases in resistance frequency. Resistance monitoring examples include Ostrinia nubilalis (Hübner) in the United States (6 yr), P. gossypiella in Arizona (5 yr), H. armigera in northern China (3 yr), and Helicoverpa zea (Boddie) in North Carolina (2 yr). Key factors delaying resistance to Bt crops are probably refuges of non-Bt host plants that enable survival of susceptible pests, low initial resistance allele frequencies, recessive inheritance of resistance to Bt crops, costs associated with resistance that reduce fitness of resistant individuals relative to susceptible individuals on non-Bt hosts ("fitness costs"), and disadvantages suffered by resistant strains on Bt hosts relative to their performance on non-Bt hosts ("incomplete resistance"). The relative importance of these factors varies among pest-Bt crop systems, and violations of key assumptions of the refuge strategy (low resistance allele frequency and recessive inheritance) may occur in some cases. The success of Bt crops exceeds expectations of many, but does not preclude resistance problems in the future.     

Comparing the refuge strategy for managing the evolution of insect resistance under different reproductive strategies

Genetically modified (GM) crops are used extensively worldwide to control diploid agricultural insect pests that reproduce sexually. However, future GM crops will likely soon target haplodiploid and parthenogenetic insects. As rapid pest adaptation could compromise these novel crops, strategies to manage resistance in haplodiploid and parthenogenetic pests are urgently needed. Here, we developed models to characterize factors that could delay or prevent the evolution of resistance to GM crops in diploid, haplodiploid, and parthenogenetic insect pests. The standard strategy for managing resistance in diploid pests relies on refuges of non-GM host plants and GM crops that produce high toxin concentrations. Although the tenets of the standard refuge strategy apply to all pests, this strategy does not greatly delay the evolution of resistance in haplodiploid or parthenogenetic pests. Two additional factors are needed to effectively delay or prevent the evolution of resistance in such pests, large recessive or smaller non-recessive fitness costs must reduce the fitness of resistance individuals in refuges (and ideally also on GM crops), and resistant individuals must have lower fitness on GM compared to non-GM crops (incomplete resistance). Recent research indicates that the magnitude and dominance of fitness costs could be increased by using specific host–plants, natural enemies, or pathogens. Furthermore, incomplete resistance could be enhanced by engineering desirable traits into novel GM crops. Thus, the sustainability of GM crops that target haplodiploid or parthenogenetic pests will require careful consideration of the effects of reproductive mode, fitness costs, and incomplete resistance.     

The cost of resistance to Bacillus thuringiensis varies with the host plant of Trichoplusia ni

Selection for resistance to insecticides, diseases and parasitoids is assumed to be costly and often requires tradeoffs with reproductive fitness. The costs of resistance, however, are often difficult to measure. Cabbage looper, Trichoplusia ni, a generalist Lepidopteran herbivore, has become highly resistant following the extensive use of the microbial insecticide, Bacillus thuringiensis kurstaki (Bt) in vegetable greenhouses. We compared the growth rate, pupal size and survival of resistant, susceptible and hybrid T. ni larvae fed on tomato, bell pepper and cucumber. Performance was best on cucumber and worst on pepper, and the magnitude of fitness costs associated with Bt resistance increased with declining host plant suitability. This supports the hypothesis that in this system, resistance costs are condition dependent and are greatest in the most stressful environment. Management strategies that rely on the presence of fitness costs to reduce the frequency of resistance genes must consider this variation and should be more successful on crops that are less suitable food plants. In general, condition dependence should be considered in studies designed to measure the costs of resistance.     

Refuges in reverse: the spread of Bacillus thuringiensis resistance to unselected greenhouse populations of cabbage loopers Trichoplusia ni

1 The dispersal of susceptible insects between refuges and Bacillus thuringiensis (Bt) treated fields is the key to resistance management of Bt crops. Here we describe the opposite situation; the movement of Bt resistant Trichoplusia ni moths from over‐wintered, greenhouse populations in British Columbia (BC) exposed to high Bt use to neighbouring greenhouses where Bt sprays have not been used. 2 The spread of Bt resistance to non‐selected populations of T. ni, and the resulting increase in resistance, indicates a surprising level of dispersal of resistant moths among greenhouses even in the face of fitness costs. 3 Field populations of T. ni in BC are seasonal migrants from regions of California where Bt cotton is grown. In 2006, field populations surveyed along the migration path from California through Oregon were highly susceptible to Bt insecticides and, thus, showed no indication of selection for resistance among these source populations. 4 The arrival of the immigrant moths provides a potential source of susceptible individuals to dilute the levels of resistance in greenhouse populations in BC later in the summer, but this has not occurred. Thus, field populations in BC do not appear to serve as refuges to combat Bt resistance in greenhouse populations.     

Evolution of resistance to transgenic crops: interactions between insect movement and field distribution

The refuge strategy is designed to delay evolution of pest resistance to transgenic crops producing Bacillus thuringiensis Berliner (Bt) toxins. Movement of insects between Bt crops and refuges of non-Bt crops is essential for the refuge strategy because it increases chances that resistant adults mate with susceptible adults from refuges. Conclusions about optimal levels of movement for delaying resistance are not consistent among previous modeling studies. To clarify the effects of movement on resistance evolution, we analyzed simulations of a spatially explicit model based partly on the interaction of pink bollworm, Pectinophora gossypiella (Saunders), with Bt cotton. We examined resistance evolution as a function of insect movement under 12 sets of assumptions about the relative abundance of Bt cotton (50 and 75%), temporal distribution of Bt cotton and refuge fields (fixed, partial rotation, and full rotation), and spatial distribution of fields (random and uniform). The results show that interactions among the relative abundance and distribution of refuges and Bt cotton fields can alter the effects of movement on resistance evolution. The results also suggest that differences in conclusions among previous studies can be explained by differences in assumptions about the relative abundance and distribution of refuges and Bt crop fields. With fixed field locations and all Bt cotton fields adjacent to at least one refuge, resistance evolved slowest with low movement. However, low movement and fixed field locations favored rapid resistance evolution when some Bt crop fields were isolated from refuges. When refuges and Bt cotton fields were rotated to the opposite crop type each year, resistance evolved fastest with low movement. Nonrecessive inheritance of resistance caused rapid resistanceevolution regardless of movement rate. Confirming previous reports, results described here show that resistance can be delayed effectively by fixing field locations and distributing refuges uniformly to ensure that Bt crop fields are not isolated from refuges. However, rotating fields provided better insect control and reduced the need for insecticide sprays.     

Modeling evolution of resistance to pyriproxyfen by the sweetpotato whitefly (Homoptera: Aleyrodidae)

We used computer simulations to examine evolution of resistance to the insect growth regulator (IGR) pyriproxyfen by the sweetpotato whitefly, Bemisia tabaci (Gennadius), biotype B [=Bemisia argentifolii (Bellows & Perring)]. Consistent with trends seen in cotton (Gossipyium spp.) fields in Arizona and Israel, results suggest that evolution of resistance to pyriproxyfen may occur rapidly in this haplodiploid insect. Similar to results from models of diploid insects, resistance evolved faster with increases in toxin concentration, dominance of resistance in females, the initial frequency of the resistance allele, and the proportion of the region treated with pyriproxyfen. Resistance was delayed by fitness costs associated with resistance. Movement between treated and untreated cotton fields had little effect, probably because untreated cotton leaves provided internal refuges in treated fields and whiteflies were controlled with other insecticides in external refuges. Resistance evolved faster when susceptibility to pyriproxyfen was greater in susceptible males than susceptible females. In contrast, resistance evolved slower when susceptibility to pyriproxyfen was greater in resistant males than resistant females. Results suggest that growers may be able to prolong the usefulness of pyriproxyfen by applying lower toxin concentrations and promoting susceptible populations in refuges.     

Delaying evolution of insect resistance to transgenic crops by decreasing dominance and heritability

The refuge strategy is used widely for delaying evolution of insect resistance to transgenic crops that produce Bacillus thuringiensis (Bt) toxins. Farmers grow refuges of host plants that do not produce Bt toxins to promote survival of susceptible pests. Many modelling studies predict that refuges will delay resistance longest if alleles conferring resistance are rare, most resistant adults mate with susceptible adults, and Bt plants have sufficiently high toxin concentration to kill heterozygous progeny from such matings. In contrast, based on their model of the cotton pest Heliothis virescens, Vacher et al. (Journal of Evolutionary Biology, 16, 2003, 378) concluded that low rather than high toxin doses would delay resistance most effectively. We demonstrate here that their conclusion arises from invalid assumptions about larval concentration-mortality responses and dominance of resistance. Incorporation of bioassay data from H. virescens and another key cotton pest (Pectinophora gossypiella) into a population genetic model shows that toxin concentrations high enough to kill all or nearly all heterozygotes should delay resistance longer than lower concentrations.     

Effects of entomopathogenic nematodes on evolution of pink bollworm resistance to Bacillus thuringiensis toxin Cry1Ac

The evolution of resistance by pests can reduce the efficacy of transgenic crops that produce insecticidal toxins from Bacillus thuringiensis (Bt). However, fitness costs may act to delay pest resistance to Bt toxins. Meta-analysis of results from four previous studies revealed that the entomopathogenic nematode Steinernema riobrave (Rhabditida: Steinernematidae) imposed a 20% fitness cost for larvae of pink bollworm, Pectinophora gossypiella (Saunders) (Lepidoptera: Gelechiidae), that were homozygous for resistance to Bt toxin Cry1Ac, but no significant fitness cost was detected for heterozygotes. We conducted greenhouse and laboratory selection experiments to determine whether S. riobrave would delay the evolution of pink bollworm resistance to Cry1Ac. We mimicked the high dose/refuge scenario in the greenhouse with Bt cotton (Gossypium hirsutum L.) plants and refuges of non-Bt cotton plants, and in the laboratory with diet containing Cry1Ac and refuges of untreated diet. In both experiments, half of the replicates were exposed to S. riobrave and half were not. In the greenhouse, S. riobrave did not delay resistance. In the laboratory, S. riobrave delayed resistance after two generations but not after four generations. Simulation modeling showed that an initial resistance allele frequency > 0.015 and population bottlenecks can diminish or eliminate the resistance-delaying effects of fitness costs. We hypothesize that these factors may have reduced the resistance-delaying effects of S. riobrave in the selection experiments. The experimental and modeling results suggest that entomopathogenic nematodes could slow the evolution of pest resistance to Bt crops, but only under some conditions.     

Early detection of field-evolved resistance to Bt cotton in China: cotton bollworm and pink bollworm

Transgenic crops producing Bacillus thuringiensis (Bt) toxins kill some major insect pests, but pests can evolve resistance and thereby reduce the effectiveness of such Bt crops. The main approach for slowing pest adaptation to Bt crops uses non-Bt host plants as "refuges" to increase survival of susceptible pests. To delay evolution of pest resistance to cotton producing Bt toxin Cry1Ac, several countries have required refuges of non-Bt cotton, while farmers in China have relied on "natural" refuges of non-Bt host plants other than cotton. This strategy is designed for cotton bollworm (Helicoverpa armigera), which attacks many crops and is the primary target of Bt cotton in China, but it does not apply to pink bollworm (Pectinophora gossypiella), which feeds almost entirely on cotton in China. Here we review evidence of field-evolved resistance to Cry1Ac by cotton bollworm in northern China and by pink bollworm in the Yangtze River Valley of China. For both pests, results of laboratory diet bioassays reveal significantly decreased susceptibility of field populations to Cry1Ac, yet field control failures of Bt cotton have not been reported. The early detection of resistance summarized here may spur countermeasures such as planting Bt cotton that produces two or more distinct toxins, increased planting of non-Bt cotton, and integration of other management tactics together with Bt cotton.     

Insect resistance to Bt crops: evidence versus theory

Evolution of insect resistance threatens the continued success of transgenic crops producing Bacillus thuringiensis (Bt) toxins that kill pests. The approach used most widely to delay insect resistance to Bt crops is the refuge strategy, which requires refuges of host plants without Bt toxins near Bt crops to promote survival of susceptible pests. However, large-scale tests of the refuge strategy have been problematic. Analysis of more than a decade of global monitoring data reveals that the frequency of resistance alleles has increased substantially in some field populations of Helicoverpa zea, but not in five other major pests in Australia, China, Spain and the United States. The resistance of H. zea to Bt toxin Cry1Ac in transgenic cotton has not caused widespread crop failures, in part because other tactics augment control of this pest. The field outcomes documented with monitoring data are consistent with the theory underlying the refuge strategy, suggesting that refuges have helped to delay resistance.     

Greenhouse tests on resistance management of Bt transgenic plants using refuge strategies

Experimental evaluation of the effectiveness of resistance management tactics is vital to help provide guidelines for the deployment of transgenic insecticidal crops. Transgenic broccoli expressing a Cry1Ac gene of Bacillus thuringiensis (Bt) and the diamondback moth, Plutella xylostella (L.), were used in greenhouse tests to evaluate the influence of size and placement of nontransgenic refuge plants on changes in resistance allele frequency and pest population growth. In the first test with an initial Cry1Ac-resistance (R) allele frequency of 0.007, P. xylostella were introduced into cages with the following treatments: 0, 3.3, 10, 20, and 100% refuge plants. Results after four generations showed that resistance could be delayed by increasing the proportion of refuge plants in the cage. Population growth was also influenced by refuge size with the highest populations occurring in treatments that had either no refuge plants or all refuge plants. In the second test, we evaluated the effect of refuge placement by comparing 20% separate and 20% mixed refuges. P. xylostella with an initial frequency of resistant alleles at 0.0125 were introduced into cages and allowed to cycle; later generations were evaluated for resistance and population growth. Separating the refuge had a pronounced effect on delaying resistance and slowing establishment of resistant larvae on Bt plants. Combining information from both trials, we found a strong negative correlation between the number of larvae on Bt plants and the mortality of the population in leaf dip bioassays. Results from larval movement studies showed that separate refuges delayed resistance better than mixed refuges because they conserved relatively more susceptible alleles than R alleles and did not increase the effective dominance of resistance.     

Suppressing resistance to Bt cotton with sterile insect releases

Genetically engineered crops that produce insecticidal toxins from Bacillus thuringiensis (Bt) are grown widely for pest control. However, insect adaptation can reduce the toxins' efficacy. The predominant strategy for delaying pest resistance to Bt crops requires refuges of non-Bt host plants to provide susceptible insects to mate with resistant insects. Variable farmer compliance is one of the limitations of this approach. Here we report the benefits of an alternative strategy where sterile insects are released to mate with resistant insects and refuges are scarce or absent. Computer simulations show that this approach works in principle against pests with recessive or dominant inheritance of resistance. During a large-scale, four-year field deployment of this strategy in Arizona, resistance of pink bollworm (Pectinophora gossypiella) to Bt cotton did not increase. A multitactic eradication program that included the release of sterile moths reduced pink bollworm abundance by >99%, while eliminating insecticide sprays against this key invasive pest.     

Genetic Resistance to Bacillus thuringiensis Alters Feeding Behaviour in the Cabbage Looper,Trichoplusia ni

Evolved resistance to xenobiotics and parasites is often associated with fitness costs when the selection pressure is absent. Resistance to the widely used microbial insecticide Bacillus thuringiensis (Bt) has evolved in several insect species through the modification of insect midgut binding sites for Bt toxins, and reports of costs associated with Bt resistance are common. Studies on the costs of Bt-resistance restrict the insect to a single artificial diet or host-plant. However, it is well documented that insects can self-select appropriate proportions of multiple nutritionally unbalanced foods to optimize life-history traits. Therefore, we examined whether Bt-resistant and susceptible cabbage loopers Trichoplusia ni differed in their nutrient intake and fitness costs when they were allowed to compose their own protein:carbohydrate diet. We found that Bt-resistant T. ni composed a higher ratio of protein to carbohydrate than susceptible T. ni. Bt-resistant males exhibited no fitness cost, while the fitness cost (reduced pupal weight) was present in resistant females. The absence of the fitness cost in resistant males was associated with increased carbohydrate consumption compared to females. We demonstrate a sex difference in a fitness cost and a new behavioural outcome associated with Bt resistance.     

Fitness costs and maternal effects associated with resistance to transgenic cotton in the pink bollworm (Lepidoptera: Gelechiidae).

Transgenic cotton producing a Bacillus thuringiensis (Bt) toxin is widely used for controlling the pink bollworm, Perctinophora gossypiella (Saunders). We compared performance of pink bollworm strains resistant to Bt cotton with performance of their susceptible counterparts on non-Bt cotton. We found fitness costs that reduced survival on non-Bt cotton by an average of 51.5% in two resistant strains relative to the susceptible strains. The survival cost was recessive in one set of crosses between a resistant strain and the susceptible strain from which it was derived. However, crosses involving an unrelated resistant and susceptible strain indicated that the survival cost could be dominant. Development time on non-Bt cotton did not differ between the two related resistant and susceptible strains. A slight recessive cost affecting development time was suggested by comparison of the unrelated resistant and susceptible strains. Maternal effects transmitted by parents that had eaten Bt-treated artificial diet as larvae had negative effects on embryogenesis, adult fertility, or both, and reduced the ability of neonates to enter cotton bolls. These results provide further evidence that fitness costs associated with the evolution of resistance to Bt cotton are substantial in the pink bollworm.     

Early warning of cotton bollworm resistance associated with intensive planting of Bt cotton in China

Transgenic crops producing Bacillus thuringiensis (Bt) toxins kill some key insect pests, but evolution of resistance by pests can reduce their efficacy. The predominant strategy for delaying pest resistance to Bt crops requires refuges of non-Bt host plants to promote survival of susceptible pests. To delay pest resistance to transgenic cotton producing Bt toxin Cry1Ac, farmers in the United States and Australia planted refuges of non-Bt cotton, while farmers in China have relied on "natural" refuges of non-Bt host plants other than cotton. Here we report data from a 2010 survey showing field-evolved resistance to Cry1Ac of the major target pest, cotton bollworm (Helicoverpa armigera), in northern China. Laboratory bioassay results show that susceptibility to Cry1Ac was significantly lower in 13 field populations from northern China, where Bt cotton has been planted intensively, than in two populations from sites in northwestern China where exposure to Bt cotton has been limited. Susceptibility to Bt toxin Cry2Ab did not differ between northern and northwestern China, demonstrating that resistance to Cry1Ac did not cause cross-resistance to Cry2Ab, and implying that resistance to Cry1Ac in northern China is a specific adaptation caused by exposure to this toxin in Bt cotton. Despite the resistance detected in laboratory bioassays, control failures of Bt cotton have not been reported in China. This early warning may spur proactive countermeasures, including a switch to transgenic cotton producing two or more toxins distinct from Cry1A toxins.     

Cadherin-based resistance to Bacillus thuringiensis cotton in hybrid strains of pink bollworm: fitness costs and incomplete resistance

Recessive resistance to Bacillus thuringiensis (Bt) cotton, Gossypium hirsutum L., in laboratory-selected strains of pink bollworm, Pectinophora gossypiella (Saunders), is associated with three resistance alleles (r1, r2, and r3) of a cadherin gene. Previous experiments based on measurement of fitness components in Bt-resistant and Bt-susceptible strains revealed that fitness costs and incomplete resistance are associated with resistance. Here, we used two hybrid strains of pink bollworm, each containing a mixture of susceptible and resistant individuals, and polymerase chain reaction (PCR) amplifications to test the association between cadherin genotype and fitness components for individuals sharing a common genetic background. All survivors on Bt cotton had two r alleles, confirming that recessive cadherin alleles are tightly linked with resistance to Bt cotton. On non-Bt cotton, significantly greater developmental time for rr than ss larvae indicated a recessive fitness cost, but costs did not affect survival or pupal weight. Incomplete resistance was manifested as longer developmental time, lower survival, and smaller pupal weight in rr individuals developing on Bt cotton compared with non-Bt cotton. As in previous experiments, no significant variation in performance on Bt cotton was detected among rr genotypes. However, a meta-analysis of data from seven experiments revealed that survival on Bt cotton relative to non-Bt cotton was lower in r2r3 and higher in r1r2 compared with the other rr genotypes. Assessment of fitness components associated with cadherin genotypes in hybrid strains of pink bollworm confirms that recessive resistance to Bt cotton is associated with recessive fitness costs and incomplete resistance.