Economic analysis of dynamic management strategies utilizing. Transgenic corn for control of western corn rootworm (Coleoptera: Chrysomelidae)

We studied management strategies for western corn rootworm, Diabrotica virgifera virgifera LeConte, using transgenic corn, Zea mays L., from both a biological and an economic perspective. In areas with and without populations adapted to a 2-yr rotation of corn and soybean (rotation-resistant), the standard management strategy was to plant 80% of a cornfield (rotated and continuous) to a transgenic cultivar each year. In each area, we also studied dynamic management strategies where the proportion of transgenic corn increased over time in a region. We also analyzed management strategies for a single field that is the first to adopt transgenic corn within a larger unmanaged region. In all areas, increasing the expression of the toxin in the plant increased economic returns. In areas without rotation-resistance, planting 80% transgenic corn in the continuous cornfield each year generated the greatest returns with a medium toxin dose or greater. In areas with alleles for rotation-resistance at low initial levels, a 2-yr rotation of nontransgenic corn and soybean, Glycine max (L.) Merr., may be the most economical strategy if resistance to crop rotation is recessive. If resistance to crop rotation is additive or dominant, planting transgenic corn in the rotated cornfield was the most effective strategy. In areas where rotation-resistance is already a severe problem, planting transgenic corn in the rotated cornfield each year was always the most economical strategy. In some cases the strategies that increased the proportion of transgenic corn in the region over time increased returns compared with the standard strategies. With these strategies the evolution of resistance to crop rotation occurred more rapidly but resistance to transgenic corn was delayed compared with the standard management strategy. In areas not managed by a regional norm, increasing the proportion of transgenic corn and increasing toxin dose in the managed field generally increased returns. In a sensitivity analysis, among the parameters investigated, only density-dependent survival affected the results.     

Effect of extended diapause on evolution of resistance to transgenic Bacillus thuringiensis corn by northern corn rootworm (Coleoptera: Chrysomelidae)

We develop a population genetics model for the northern corn rootworm, Diabrotica barberi Smith & Lawrence, to examine the effect of extended diapause on the evolution of resistance to transgenic Bacillus thuringiensis (Bt) corn, Zea mays L. We model conditions found in the center of the extended diapause problem along the Minnesota-South Dakota-Iowa borders. The proportion of resistance alleles in eggs oviposited after 15 simulated years is used to measure the evolution of resistance. Sensitivity analysis indicates that although population genetics parameters (fecundity, initial egg density, density-dependent larval survival, random mating, insecticide mortality, and gene expression) affect the evolution of resistance, product characteristics (e.g., Bt toxin dose) and farmer management practices (e.g., insecticide use on refuge corn and rotation pattern) generally have a larger impact on the development of resistance. Exceptions to this generalization exist: 1) if the resistance allele is dominant, resistance evolves quickly; 2) the level of random mating is an important determinant of how quickly resistance evolves for a theoretical high dose product; and 3) small differences in insecticide mortality imply large differences in resistance for medium- and low-dose products with high levels of Bt corn adoption and a predominance of 1- and 2-yr corn rotations. When extended diapause spreads into a new area, it typically reduces resistance to Bt corn, assuming Bt corn is used only on continuous corn. In the study region where extended diapause already exists, increasing extended diapause (increasing hatch rates after two or three winters while holding total hatch constant), tends to increase resistance because the resistance increasing effect of the hatch rate after two winters dominates the resistance decreasing effect of the hatch rate after three winters. However, this is not always the case, because combinations of rotation pattern, toxin dose, and soil insecticide use exist for which the net effect of extended diapause decreases resistance. Results are interpreted as a combination of two offsetting effects. First, extended diapause injects older alleles with lower resistance allele frequencies into the breeding population, which slows resistance. Second, extended diapause speeds the population's recovery from perturbations (reduces the undercompensating density dependence of population dynamics), which accelerates resistance.     

Modeling the dynamics of adaptation to transgenic corn by western corn rootworm (Coleoptera: Chrysomelidae)

A simulation model of the population dynamics and genetics of the western corn rootworm, Diabrotica virgifera virgifera LeConte, was created for a landscape of corn, soybean, and other crops. Although the model was created to study a 2-locus problem for beetles having genes for resistance to both crop rotation and transgenic corn, during this first phase of the project, the model was simulated to evaluate only resistance management plans for transgenic corn. Allele expression in the rootworm and toxin dose in the corn plant were the two most important factors affecting resistance development. A dominant resistance allele allowed quick evolution of resistance to transgenic corn, whereas a recessive allele delayed resistance >99 yr. With high dosages of toxin and additive expression, the time required to reach 3% resistance allele frequency ranged from 13 to >99 yr. With additive expression, lower dosages permitted the resistant allele frequency to reach 3% in 2-9 yr with refuges occupying 5-30% of the land. The results were sensitive to delays in emergence by susceptible adults and configuration of the refuge (row strips versus blocks).     

Western corn rootworm (Coleoptera: Chrysomelidae) dispersal and adaptation to single-toxin transgenic corn deployed with block or blended refuge

A simulation model of the temporal and spatial dynamics and population genetics of western corn rootworm, Diabrotica virgifera virgifera LeConte, was created to evaluate the use of block refuges and seed blends in the management of resistance to transgenic insecticidal corn (Zea mays L.). This Bt corn expresses one transgenic corn event, DAS-59122-7, that produces a binary insecticidal protein toxin (Cry34Ab1/Cry35Ab1) and provides host-plant resistance. The model incorporates the latest information about larval and adult behavior. Results of this modeling effort indicate that the seed-blend scenarios in many cases produced equal or greater durability than block refuges that were relocated each year. Resistance evolved in the most likely scenarios in 10-16 yr. Our standard analysis presumed complete adoption of 59122 corn by all farmers in our hypothetical region, no crop rotation, and 100% compliance with Insect Resistant Management (IRM) regulations. As compliance levels declined, resistance evolved faster when block refuges were deployed. Seed treatments that killed the pest when applied to all seeds in a seed blend or just to seeds in Bt corn blocks delayed evolution of resistance. Greater control of the pest population by the seed treatment facilitated longer durability of the transgenic trait. Therefore, data support the concept that pyramiding a transgenic insecticidal trait with a highly efficacious insecticidal seed treatment can delay evolution of resistance.     

Seeds of change: corn seed mixtures for resistance management and integrated pest management

The use of mixtures of transgenic insecticidal seed and nontransgenic seed to provide an in-field refuge for susceptible insects in insect-resistance-management (IRM) plans has been considered for at least two decades. However, the U.S. Environmental Protection Agency has only recently authorized the practice. This commentary explores issues that regulators, industry, and other stakeholders should consider as the use of biotechnology increases and seed mixtures are implemented as a major tactic for IRM. We discuss how block refuges and seed mixtures in transgenic insecticidal corn, Zea mays L., production will influence integrated pest management (IPM) and the evolution of pest resistance. We conclude that seed mixtures will make pest monitoring more difficult and that seed mixtures may make IRM riskier because of larval behavior and greater adoption of insecticidal corn. Conversely, block refuges present a different suite of risks because of adult pest behavior and the lower compliance with IRM rules expected from farmers. It is likely that secondary pests not targeted by the insecticidal corn as well as natural enemies will respond differently to block refuges and seed mixtures.     

Modeling the development of resistance by stalk-boring lepidopteran insects (Crambidae) in areas with transgenic corn and frequent insecticide use

We simulated the population dynamics and population genetics of two bivoltine species of corn borers, the European corn borer, Ostrinia nubilalis (Hübner), and the southwestern corn borer, Diatraea grandiosella Dyar, in a hypothetical region of irrigated transgenic and nontransgenic corn where insecticide was applied only to the nontransgenic refuge crop. Over the 100-yr time horizon, resistance developed quickly in both species and to both transgenic corn and the insecticide when the allele for resistance to the respective toxin was dominant. When the allele for transgenic resistance was not dominant and the refuge location was constant over the time horizon, spraying the refuge to control southwestern corn borer had no effect on how quickly resistance to the transgenic corn developed. In contrast, the European corn borer developed resistance to transgenic corn much sooner when the refuge was sprayed once per year, and the time to 3% resistance allele frequency decreased as efficacy of the insecticide increased. Only when the refuge was treated less than once every 5 yr (10 generations) did the frequency of application decline enough to permit resistance management for the European corn borer to approximate the effectiveness of an unsprayed refuge. A consistently sprayed refuge <40% of the corn acreage was an inadequate resistance management strategy for the European corn borer even when a low efficacy insecticide (70% mortality) was used. When assumptions about European corn borer adult behavior were changed and the adults behaved similarly to adult southwestern corn borer, the development of resistance to the transgenic crop was slowed significantly.     

Economics versus alleles: balancing integrated pest management and insect resistance management for rotation-resistant western corn rootworm (Coleoptera: Chrysomelidae)

Western corn rootworm, Diabrotica virgifera virgifera LeConte, has overcome crop rotation in several areas of the central United States. We expanded a simple model of adult behavior and population genetics to explain how rotation resistance may have developed and to study ways to manage the western corn rootworm in a landscape of corn, soybean, and winter wheat where evolution of resistance may occur. We modeled six alternative management strategies over a 15-yr time horizon, as well as a strategy involving a 2-yr rotation of corn and soybean in 85% of the landscape, to investigate their effectiveness from both a biological and economic perspective. Generally, resistance to crop rotation evolves in fewer than 15 yr, and the rate of evolution increases as the level of rotated landscape (selection pressure) increases. When resistance is recessive, all six alternative strategies were effective at preventing evolution of rotation resistance. The two most successful strategies were the use of transgenic rotated corn in a 2-yr rotation and a 3-yr rotation of corn, soybean, and wheat with unattractive wheat (for oviposition) preceding corn. Results were most sensitive to increases in the initial allele frequency and modifications of the density-dependent survival function. Economically, three alternative strategies were robust solutions to the problem, if technology fees were not too high. Repellant soybean, attractive rotated corn, and transgenic rotated corn, all in 2-yr rotations, were economically valuable approaches. However, even the currently common 2-yr rotation was economical when resistance was recessive and the actual costs of resistance would not be paid until far in the future.     

Inheritance of Cry1F resistance in laboratory-selected European corn borer and its survival on transgenic corn expressing the Cry1F toxin

A major assumption of the high-dose/refuge strategy proposed for insect resistance management strategies for transgenic crop plants that express toxins from Bacillus thuringiensis is that resistance traits that evolve in pest species will be recessive. The inheritance of Cry1F resistance and larval survival on commercially available Cry1F corn hybrids were determined in a laboratory-selected strain of European corn borer, Ostrinia nubilalis (Hübner), displaying more than 3000-fold resistance to Cry1F. Concentration-response bioassays of reciprocal parental crosses indicated that the resistance is autosomal and recessive. Bioassays of the backcross of the F1 generation with the selected strain were consistent with the hypothesis that a single locus, or a set of tightly linked loci, is responsible for the resistance. Greenhouse experiments with Cry1F-expressing corn hybrids indicated that some resistant larvae survived the high dose of toxin delivered by Cry1F-expressing plants although F1 progeny of susceptible by resistant crosses had fitness close to zero. These results provide the first direct evidence that the high dose/refuge strategy currently in place to manage resistance in Cry1F-expressing corn is appropriate.     

Analysis of the dynamics of adaptation to transgenic corn and crop rotation by western corn rootworm (Coleoptera: Chrysomelidae) using a daily time-step model

Western corn rootworm, Diabrotica virgifera virgifera LeConte, has overcome crop rotation in several areas of the north central United States. The effectiveness of crop rotation for management of corn rootworm has begun to fail in many areas of the midwestern United States, thus new management strategies need to be developed to control rotation-resistant populations. Transgenic corn, Zea mays L., effective against western corn rootworm, may be the most effective new technology for control of this pest in areas with or without populations adapted to crop rotation. We expanded a simulation model of the population dynamics and genetics of the western corn rootworm for a landscape of corn; soybean, Glycine max (L.); and other crops to study the simultaneous development of resistance to both crop rotation and transgenic corn. Results indicate that planting transgenic corn to first-year cornfields is a robust strategy to prevent resistance to both crop rotation and transgenic corn in areas where rotation-resistant populations are currently a problem or may be a problem in the future. In these areas, planting transgenic corn only in continuous cornfields is not an effective strategy to prevent resistance to either trait. In areas without rotation-resistant populations, gene expression of the allele for resistance to transgenic corn, R, is the most important factor affecting the evolution of resistance. If R is recessive, resistance can be delayed longer than 15 yr. If R is dominant, resistance may be difficult to prevent. In a sensitivity analysis, results indicate that density dependence, rotational level in the landscape, and initial allele frequency are the three most important factors affecting the results.     

Using a generational time-step model to simulate dynamics of adaptation to transgenic corn and crop rotation by western corn rootworm (Coleoptera: Chrysomelidae)

We expanded a simulation model of the population dynamics and genetics of the western corn rootworm for a landscape of corn, soybean, and other crops to study the simultaneous development of resistance to both crop rotation and transgenic corn. Transgenic corn effective against corn rootworm was recently approved in 2003 and may be a very effective new technology for control of western corn rootworm in areas with or without the rotation-resistant variant. In simulations of areas with rotation-resistant populations, planting transgenic corn to only rotated cornfields was a robust strategy to prevent resistance to both traits. In these areas, planting transgenic corn to only continuous fields was not an effective strategy for preventing adaptation to crop rotation or transgenic corn. In areas without rotation-resistant phenotypes, gene expression of the allele for resistance to transgenic corn was the most important factor affecting the development of resistance to transgenic corn. If the allele for resistance to transgenic corn is recessive, resistance can be delayed longer than 15 yr, but if the resistant allele is dominant then resistance usually developed within 15 yr. In a sensitivity analysis, among the parameters investigated, initial allele frequency and density dependence were the two most important factors affecting the evolution of resistance. We compared the results of this simulation model with a more complicated model and results between the two were similar. This indicates that results from a simpler model with a generational time-step can compare favorably with a more complex model with a daily time-step.     

Evaluation of oviposition deterrence in management of resistance to transgenic corn by European corn borer (Lepidoptera: Crambidae)

We modified an existing model for European corn borer, Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae), population dynamics and genetics to evaluate the effectiveness of oviposition deterrence in transgenic fields for resistance management. We simulated two types of deterrence: one type has females reducing their oviposition because of lost opportunities to lay eggs (eggs lost), and the other type has the deterred females moving to the refuge to lay eggs. Oviposition deterrence was clearly effective in extending the time to resistance to transgenic corn (R allele) in the European corn borer, particularly when 80% or more of the eggs were deterred from being oviposited on the transgenic plants. With 90% of eggs deterred, the time required to reach 50% R-allele frequency increases 3.7- to 5.5-fold compared with the no-deterrence scenario. The time to 50% R-allele frequency was similar for the two types of simulated deterrence, but the densities of the European corn borer were 100-fold higher when the deterred females oviposited in the refuge. The Y allele for insensitivity or resistance to deterrence never reached 50% within the 50-yr time line for these simulations except when the R allele was dominant and the Y allele was not recessive. The time to 50% Y-allele frequency was 33 and 26 yr when the Y allele was additive or dominant, respectively, when 50% of the eggs were deterred, but the time decreased to 18 and 16 yr when 90% of the eggs were deterred. The effectiveness of oviposition deterrence on time to resistance to transgenic insecticidal plants was not changed much when we altered our assumptions about behavior in a sensitivity analysis.     

Seed mixtures as a resistance management strategy for European corn borers (Lepidoptera: Crambidae) infesting transgenic corn expressing Cry1Ab protein

Dispersal of neonate European corn borers, Ostrinia nubilalis (Hübner), in seed mixtures of transgenic corn expressing Cry1Ab protein (Bt+) and nontransgenic corn (Bt-) was evaluated in a 2-yr field study. The main objective was to determine if larval dispersal limits the effectiveness of seed mixtures as a resistance management strategy. Mixtures evaluated included (1) all Bt+ plants, (2) every fifth plant Bt- with remaining plants Bt+, (3) every fifth plant Bt+ with remaining plants Bt-, and (4) all Bt- plants. The transformation events MON 802 (B73 BC1F2 x Mol7) and MON 810 (B73 BC1F1 x Mo17), which express the Cry1Ab endotoxin isolated from Bacillus thuringiensis subsp. kurstaki, were used as the sources of Bt+ seed in 1994 and 1995, respectively (YieldGard, Monsanto, St. Louis, MO). At corn growth stage V6-V8, subplots within each mixture (15-20 plants each) were infested so that every fifth plant in mixtures 1 and 4, every Bt- plant in mixture 2, and every Bt+ plant in mixture 3 received two egg masses. Larval sampling over a 21-d period indicated increased neonate dispersal off of Bt+ plants, reduced survival of larvae that dispersed from Bt+ plants to Bt- plants, and a low incidence of late-instar movement from Bt- plants to Bt+ plants. Computer simulations based on mortality and dispersal estimates from this study indicate that seed mixtures will delay the evolution of resistant European corn borer populations compared with uniform planting of transgenic corn. However, resistant European corn borer populations likely will develop faster in seed mixes compared with separate plantings of Bt and non-Bt corn.     

Reversing insect adaptation to transgenic insecticidal plants.

The refuge-high-dose strategy for delaying insect adaptation to transgenic plants produces non-transgenic plants that enable survival of susceptible individuals. Previous theoretical work has suggested three requirements for success of the refuge-high-dose strategy: a low initial frequency of the resistance allele, extensive mating between resistant and susceptible adults and recessive inheritance of resistance. In order to understand an observed decrease in resistance frequency and improve the potential for managing resistance better, we used analytical and simulation models for exploring the conditions that prevent or reverse the evolution of resistance, even when resistance is not rare initially. Assuming random mating and recessive or nearly recessive inheritance of resistance, the factors favouring reversal of resistance are non-recessive costs of resistance, low initial resistance allele frequency, large refuges, incomplete resistance and density-independent population growth in refuges.     

Delaying corn rootworm resistance to Bt corn

Transgenic crops producing Bacillus thuringiensis (Bt) toxins for insect control have been successful, but their efficacy is reduced when pests evolve resistance. To delay pest resistance to Bt crops, the U.S. Environmental Protection Agency (EPA) has required refuges of host plants that do not produce Bt toxins to promote survival of susceptible pests. Such refuges are expected to be most effective if the Bt plants deliver a dose of toxin high enough to kill nearly all hybrid progeny produced by matings between resistant and susceptible pests. In 2003, the EPA first registered corn, Zea mays L., producing a Bt toxin (Cry3Bb1) that kills western corn rootworm, Diabrotica virgifera virgifera LeConte, one of the most economically important crop pests in the United States. The EPA requires minimum refuges of 20% for Cry3Bb1 corn and 5% for corn producing two Bt toxins active against corn rootworms. We conclude that the current refuge requirements are not adequate, because Bt corn hybrids active against corn rootworms do not meet the high-dose standard, and western corn rootworm has rapidly evolved resistance to Cry3Bb1 corn in the laboratory, greenhouse, and field. Accordingly, we recommend increasing the minimum refuge for Bt corn targeting corn rootworms to 50% for plants producing one toxin active against these pests and to 20% for plants producing two toxins active against these pests. Increasing the minimum refuge percentage can help to delay pest resistance, encourage integrated pest management, and promote more sustainable crop protection.     

Modeling the impact of cross-pollination and low toxin expression in corn kernels on adaptation of European corn borer (Lepidoptera: Crambidae) to transgenic insecticidal corn

We used a mathematical model with processes reflecting larval mortality resulting from feeding on cross-pollinated ears or Bt ears of corn to analyze the risk of evolution of Cry-toxin resistance in Ostrinia nubilalis (Hübner). In the simulations, evolution of resistance was delayed equally well by both seed mixtures and blocks with the same proportion of refuge. Our results showed that Bt-pollen drift has little impact on the evolution of Bt resistance in O. nubilalis. However, low-toxin expression in ears of transgenic corn can reduce the durability of transgenic corn expressing single toxin, whereas durability of pyramided corn hybrids is not significantly reduced. The toxin-survival rate of heterozygous larvae in Bt-corn ears expressing one or two proteins has more impact on evolution of Bt resistance in O. nubilalis than the parameters related to larval movement to Bt ears or the toxin-survival rate of the homozygous susceptible larvae in Bt ears. Bt resistance evolves slower when toxin mortality is distributed across the first two larval stadia than when only the first instars are susceptible to Bt toxins. We suggest that stakeholders examine toxin-survival rates for insect pests and take into account that instars may feed on different parts of Bt corn.     

Impact of small fitness costs on pest adaptation to crop varieties with multiple toxins: a heuristic model

A deterministic two-locus model was used to examine how small fitness costs to individuals carrying resistance alleles could impact the risk of panmictic insect pest populations adapting to crop varieties that produced two distinct toxins. Parameters examined were (1) level of toxicity of each toxin, (2) initial frequencies of alleles for adaptation to the toxins, (3) percentage of population feeding on nontoxic plants, and (4) level of fitness cost associated with adaptation to each of the two toxins. Resistance to each toxin was assumed to be biochemically independent, controlled by a resistance coding allele at a single locus, and inherited as a partially recessive trait in the field. When plants are extremely toxic to the pest, effective refuge size is 10%, and there is a fitness cost to resistance alleles only when in homozygous form (5%), the pest population is never predicted to adapt to either toxin as long as the initial frequencies of the resistance alleles are below 0.05. Even if the initial frequency of the allele for adapting to one toxin is 0.95 when a two-toxin cultivar completely replaces a one-toxin cultivar, the model predicts that a low equilibrium allelic frequency will develop for both resistance alleles, as long as the frequency of the allele for adapting to the second toxin is initially 0.001 or less. If cultivars with one and two toxins are planted, the model predicts that resistance will develop. Nonrandom mating and stochastic variation within subpopulations also could lead to evolution of resistance.     

Cross-pollination of nontransgenic corn ears with transgenic Bt corn: efficacy against lepidopteran pests and implications for resistance management

The efficacy of nontransgenic sweet corn, Zea mays L., hybrids cross-pollinated by Bacillus thuringiensis (Bt) sweet corn hybrids expressing Cry1Ab toxin was evaluated in both field and laboratory studies in Minnesota in 2000. Non-Bt and Bt hybrids (maternal plants) were cross-pollinated with pollen from both non-Bt and Bt hybrids (paternal plants) to create four crosses. Subsequent crosses were evaluated for efficacy in the field against European corn borer, Ostrinia nubilalis (Hübner), and corn earworm, Helicoverpa zea (Boddie), and in laboratory bioassays against O. nubilalis. Field studies indicated that crosses with maternal Bt plants led to low levels of survival for both O. nubilalis and H. zea compared with the non-Bt x non-Bt cross. However, the cross between non-Bt ears and Bt pollen led to survival rates of 43 and 63% for O. nubilalis and H. zea larvae, respectively. This intermediate level of survival also was reflected in the number of kernels damaged. Laboratory bioassays for O. nubilalis, further confirmed field results with larval survival on kernels from the cross between non-Bt ears and Bt pollen reaching 60% compared with non-Bt crossed with non-Bt. These results suggest that non-Bt refuge plants, when planted in proximity to Bt plants, and cross-pollinated, can result in sublethal exposure of O. nubilalis and H. zea larvae to Bt and may undermine the high-dose/refuge resistance management strategy for corn hybrids expressing Cry1Ab.     

Stochastic dynamic population model for northern corn rootworm (Coleoptera: Chrysomelidae).

A stochastic dynamic population model for the complete life cycle of northern corn rootworm, Diabrotica barberi Smith & Lawrence, is described. Adult population dynamics from emergence to oviposition are based on a published single-season model for which temperature-dependent development and age-dependent advancement determine adult population dynamics and oviposition. Randomly generated daily temperatures make this model component stochastic. Stochastic hatch is 50+/-8%. A stochastic nonlinear density-dependent larval survival model is estimated using field data from artificial infestation experiments. A regional model of corn phenology is estimated to incorporate the effect of dispersal on adult mortality. Random daily weather is generated using parameters for Brookings, SD. Model performance is evaluated with deterministic simulations, which show that the population converges to zero unless adult mortality is reduced by the availability of corn pollen from the regional model of corn phenology. Stochastic model performance is evaluated with stochastic daily weather, egg hatch, and larval survival in various combinations. Sensitivity analysis is conducted to evaluate model responsiveness to each parameter. Model results are generally consistent with published data.     

A spatially explicit model simulating western corn rootworm (Coleoptera: Chrysomelidae) adaptation to insect-resistant maize

A stochastic spatially explicit computer model is described that simulates the adaptation by western corn rootworm, Diabrotica virgifera virgifera LeConte, to rootworm-resistance traits in maize. The model reflects the ecology of the rootworm in much of the corn belt of the United States. It includes functions for crop development, egg and larval mortality, adult emergence, mating, egg laying, mortality and dispersal, and alternative methods of rootworm control, to simulate the population dynamics of the rootworm. Adaptation to the resistance trait is assumed to be controlled by a monogenic diallelic locus, whereby the allele for adaptation varies from incompletely recessive to incompletely dominant, depending on the efficacy of the resistance trait. The model was used to compare the rate at which the adaptation allele spread through the population under different nonresistant maize refuge deployment scenarios, and under different levels of crop resistance. For a given refuge size, the model indicated that placing the nonresistant refuge in a block within a rootworm-resistant field would be likely to delay rootworm adaptation rather longer than planting the refuge in separate fields in varying locations. If a portion of the refuge were to be planted in the same fields or in-field blocks each year, rootworm adaptation would be delayed substantially. Rootworm adaptation rates are also predicted to be greatly affected by the level of crop resistance, because of the expectation of dependence of functional dominance on dose. If the dose of the insecticidal protein in the maize is sufficiently high to kill >90% of heterozygotes and approximately 100% of susceptible homozygotes, the trait is predicted to be much more durable than if the dose is lower. A partial sensitivity analysis showed that parameters relating to adult dispersal affected the rate of pest adaptation. Partial validation of the model was achieved by comparing output of the model with field data on population dynamics, and with field data documenting rootworm adaptation to cyclodienes and organophosphates.     

Characterizing laboratory colonies of western corn rootworm (Coleoptera: Chrysomelidae) selected for survival on maize containing event DAS-59122-7

Event DAS‐59122‐7 is a novel transgenic trait designed to protect the roots and yield potential of maize from the insect pest corn rootworm Diabrotica spp. (Col.: Chrysomelidae). The increased pest status of corn rootworm, exceptional efficacy of this trait, and anticipated increases in farm efficiency and grower and environmental safety will drive adoption of this trait. Strong grower acceptance of this trait highlights the importance of science‐based and practical resistance management strategies. A non‐diapause trait was introgressed into two laboratory colonies of Diabrotica virgifera virgifera collected from geographically distinct locations: Rochelle, IL and York, NE. Both colonies were divided and each reared on maize containing event DAS‐59122‐7 or its near isoline. Selected and unselected colonies were evaluated for phenotypic change in larval development, injury potential and survival to adulthood during 10 and 11 generations. The F1 generation of both selected colonies displayed increased larval development, survivorship and measurable, but economically insignificant increases in injury potential on DAS‐59122‐7 maize. Survival rates of 0.4 and 1.3% in F1 generations of both selected colonies corroborate field estimates of survival on DAS‐59122‐7 maize. Over later generations, total phenotypic variation declined gradually and irregularly. Despite the absence of random mating, the tolerance trait could not be fixed in either population after 10 or 11 generations of selection. An allele conferring major resistance to DAS‐59122‐7 was not identified in either selected colony. The assessment also concluded that major resistance gene(s) are rare in populations of D. v. virgifera in the United States, and that a minor trait(s) conferring a low level of survival on DAS‐59122‐7 maize was present. The tolerance trait identified in this study was considered minor with respect to its impact on DAS‐59122‐7 maize efficacy, and the role this trait may play in total effective refuge for major resistance genes with recessive inheritance is the basis of future work.