Resistance to Bt Corn by Western Corn Rootworm (Coleoptera: Chrysomelidae) in the U.S. Corn Belt

Transgenic Bt corn hybrids that produce insecticidal proteins from the bacterium Bacillus thuringiensis Berliner have become the standard insect management tactic across the U.S. Corn Belt. Widespread planting of Bt corn places intense selection pressure on target insects to develop resistance, and evolution of resistance threatens to erode benefits associated with Bt corn, such as reduced reliance on conventional insecticides. Recognizing the threat of resistance, the U.S. Environmental Protection Agency requires seed companies to include an insect resistance management (IRM) plan when registering a Bt trait. The goal of IRM plans is to delay Bt resistance in populations of target insects. One element of IRM is the presence of a non-Bt refuge to maintain Bt-susceptible individuals within a population, and growers are required to implement IRM on-farm by planting a refuge. Field-evolved resistance has not been detected for the European corn borer, Ostrinia nubilalis (Hubner), even though this species has been exposed to Bt proteins common in U.S. corn hybrids since 1996. The IRM situation is unfolding differently for Bt corn targeting the western corn rootworm, Diabrotica virgifera virgifera LeConte. In this article, we examine the scientific evidence for D. v. virgifera resistance to Bt rootworm traits and the cropping system practices that have contributed to the first reports of field-evolved resistance to a Bt toxin by D. v. virgifera. We explain why this issue has developed, and emphasize the necessity of an integrated pest management approach to address the issue

Adaptation by Western Corn Rootworm (Coleoptera: Chrysomelidae) to Bt Maize: Inheritance, Fitness Costs, and Feeding Preference

We examined inheritance of resistance, feeding behavior, and fitness costs for a laboratory-selected strain of western corn rootworm, Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae), with resistance to maize (Zea maize L.) producing the Bacillus thuringiensisBerliner (Bt) toxin Cry3Bb1. The resistant strain developed faster and had increased survival on Bt maize relative to a susceptible strain. Results from reciprocal crosses of the resistant and susceptible strains indicated that inheritance of resistance was nonrecessive. No fitness costs were associated with resistance alleles in the presence of two entomopathogenic nematode species, Steinernema carpocapsae Weiser and Heterorhabditis bacteriophora Poinar. Larval feeding studies indicated that the susceptible and resistant strains did not differ in preference for Bt and non-Bt root tissue in choice assays

Field-Evolved Resistance to Bt Maize by Western Corn Rootworm

Crops engineered to produce insecticidal toxins derived from the bacterium Bacillus thuringiensis (Bt) are planted on millions of hectares annually, reducing the use of conventional insecticides and suppressing pests. However, the evolution of resistance could cut short these benefits. A primary pest targeted by Bt maize in the United States is the western corn rootwormDiabrotica virgifera virgifera (Coleoptera: Chrysomelidae). We report that fields identified by farmers as having severe rootworm feeding injury to Bt maize contained populations of western corn rootworm that displayed significantly higher survival on Cry3Bb1 maize in laboratory bioassays than did western corn rootworm from fields not associated with such feeding injury. In all cases, fields experiencing severe rootworm feeding contained Cry3Bb1 maize. Interviews with farmers indicated that Cry3Bb1 maize had been grown in those fields for at least three consecutive years. There was a significant positive correlation between the number of years Cry3Bb1 maize had been grown in a field and the survival of rootworm populations on Cry3Bb1 maize in bioassays. However, there was no significant correlation among populations for survival on Cry34/35Ab1 maize and Cry3Bb1 maize, suggesting a lack of cross resistance between these Bt toxins. This is the first report of field-evolved resistance to a Bt toxin by the western corn rootworm and by any species of Coleoptera. Insufficient planting of refuges and non-recessive inheritance of resistance may have contributed to resistance. These results suggest that improvements in resistance management and a more integrated approach to the use of Bt crops may be necessary

Field-Based Assessment of Resistance to Bt Corn by Western Corn Rootworm (Coleoptera: Chrysomelidae)

Western corn rootworm, Diabrotica virgifera virgifera LeConte, is a serious pest of corn and is managed with corn that produces insecticidal toxins derived from the bacterium Bacillus thuringiensis (Bt). Beginning in 2009, resistance to Cry3Bb1 corn, and severe injury to Cry3Bb1 corn in the field, was observed in Iowa. However, few data exist on how Cry3Bb1-resistant western corn rootworm interact with various management practices in the field. Using a field experiment, we measured adult emergence and feeding injury to corn roots for both Cry3Bb1-resistant and Cry3Bb1-susceptible populations of western corn rootworm when tested against various Bt corn hybrids and a soil-applied insecticide. Between 2012 and 2013, we evaluated five fields that were associated with greater than one node of feeding injury to Cry3Bb1 corn by western corn rootworm (i.e., problem-field populations), and a laboratory strain that had never been exposed to Bt corn (i.e., control population). Adult emergence for western corn rootworm and root injury to corn were significantly higher in problem-field populations than control populations for both Cry3Bb1 corn and mCry3A corn. By contrast, corn with Cry34/35Ab1, either alone or pyramided with Cry3Bb1, significantly reduced adult emergence and root injury in both problem fields and control fields. In problem fields, application of the soil-applied insecticide to Cry3Bb1 corn significantly reduced root injury, but not adult emergence. Our results are discussed in terms of developing strategies for managing western corn rootworm with resistance to Cry3Bb1 and mCry3A, and delaying the additional evolution of Bt resistance by this pest

Entomopathogenic fungi in cornfields and their potential to manage larval western corn rootworm Diabrotica virgifera virgifera

Entomopathogenic ascomycete fungi are ubiquitous in soil and on phylloplanes, and are important natural enemies of many soil-borne arthropods including larval western corn rootworm, Diabrotica virgifera virgifera, which is a major pest of corn. We measured the prevalence of Beauveria bassiana andMetarhizium anisopliae sensu lato in ten cornfields in Iowa, USA by baiting with larval insects. B. bassianaand M. anisopliae s.l. were present in 60% ± 6.3% and 55% ± 6.4% of soil samples, respectively. Subsequent laboratory bioassays found that some M. anisopliae s.l. strains collected from cornfields killed a greater proportion of D.v. virgifera larvae than a standard commercial strain

Evaluation of Bt Corn and Soil-applied Insecticides for Management of Corn Rootworm Larvae

The purpose of this study was to evaluate the effectiveness of Bt corn, with and without soil insecticides, for management of corn rootworm. Evaluation of Bt hybrids included Agrisure 3000GT, Agrisure 3122 RIB, DeKalb YieldGard VT3, and Pioneer Optimum AcreMax1. Soil insecticides evaluated were Aztec 2.1G, Aztec-SB 4.67G, Capture LFR 1.5SC, Counter-SB 20G, Force 250CS, and SmartChoice-SB 5G

Effect of Bt Maize and Soil Insecticides on Yield, Injury, and Rootworm Survival: Implications for Resistance Management

A 2-yr field experiment was conducted to determine the effects on Diabrotica spp. (Coleoptera: Chrysomelidae) of an insecticidal seed treatment (Poncho 1250, (AI) /clothianidin) and a granular insecticide (Aztec 2.1G, (AI)/tebupirimphos and cyfluthrin) alone and in combination with maize producing the insectidical toxin Cry3Bb1 derived from the bacterium Bacillus thuringiensis (Bt). Yields for Bt maize plots were significantly greater than for non-Bt maize; however, insecticides did not significantly affect yield. Insecticides significantly decreased root injury in non-Bt maize plots, but there were no significant differences in root injury between Bt maize with or without either insecticide. Maize producing the Bt toxin Cry3Bb1 and the soil-applied insecticide Aztec significantly decreased survival of western corn rootworm (Diabrotica virgifera virgifera LeConte), while only Bt maize significantly decreased survival of the northern corn rootworm (Diabrotica barberi Smith & Lawrence). For both species, Bt maize and each of the insecticides delayed emergence. In the absence of density-dependent mortality, Bt maize imposed 71 and 80% reduction in survival on the western corn rootworm and the northern corn rootworm, respectively. The data from this study do not support combining insecticide with Bt maize because the addition of insecticide did not increase yield or reduce root injury for Bt maize, and the level of rootworm mortality achieved with conventional insecticide was likely too low to delay the evolution of Bt resistance. In addition, delays in emergence from Bt maize combined with insecticides could promote assortative mating among Bt-selected individuals, which may hasten resistance evolution

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 of transgenic 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 nematodeSteinernema riobrave Cabanillas, Poinar, and Raulston but no fitness costs were imposed bySteinernema carpocapsae Weiser, Steinernema sp. (ML18 strain), or Heterorhabditis sonorensis Stock, Rivera-Orduño, and Flores-Lara. In computer simulations, evolution of resistance to CrylAc 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

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 \u3e0.015 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

How Many Corn Traits Do You Need?

We seem to live in a “have it your way” world. Everything from fast food hamburgers to “designer” clothing tailored in countries some of us didn’t know existed. In contrast to this, modern, mass-production systems encourage wholesale consumption of identical products worldwide: “one-size fits all.” These contradictions also exist in the world of corn production