Estimating western corn rootworm (Coleoptera: Chrysomelidae) larval susceptibility to event DAS‐59122‐7 maize

Susceptibility of Diabrotica virgifera virgifera (LeConte) larvae to DAS‐59122‐7 maize was evaluated using a laboratory technique that measures rootworm survival to adulthood on maize seedlings. This method produces direct measures of larval susceptibility using realistic exposure to the same range of insecticidal protein concentrations found in field‐grown DAS‐59122‐7 maize roots. First, second and third instars were reared to adulthood on DAS‐59122‐7 maize seedlings or a non‐transgenic, near‐isoline maize. Data on survival, adult gender ratio, adult weight and median emergence were collected. Overall, larval susceptibility to DAS‐59122‐7 maize was lower than earlier predictions ( Storer et al. 2006 ). Neonate survival on DAS‐59122‐7 maize was approximately 33% of isoline survival after 17 days, and the same 33% recovered and developed to adulthood when the isoline maize was substituted. Survival rate on DAS‐59122‐7 maize increased with instar. The mean survivorship was 0.5%, 26% and 65% when exposure to DAS‐59122‐7 maize began at the first, second and third instars, respectively. Exposure to DAS‐59122‐7 maize led to sub‐lethal effects on adult gender ratio, weight and median emergence. These effects decreased when exposure to DAS‐59122‐7 maize began at later instars. The killing effect of DAS‐59122‐7 maize on rootworm larvae appeared to result from the combined chronic effects and absence of a suitable host as perceived by the larvae. The relevance of these data and the methodology of estimating rootworm susceptibility to plant‐incorporated protectants are discussed in the context of the US Environmental Protection Agency’s functional definition of ‘high dose’ and use of refuge for resistance management ( EPA 1998a ). Based on these results it is evident that DAS‐59122‐7 maize does not meet the functional definitions of high dose as described by EPA (1998a,b) and ILSI (1999) , and the utility of refuge, refuge size and refuge placement for delaying rootworm resistance should be further investigated.     

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.     

Development and economic evaluation of spatial sampling plans for corn rootworm Diabrotica spp. (Col., Chrysomelidae) adults

Abstract  To develop spatial sampling plans for corn rootworm ( Diabrotica spp.) adults, their spatial distributions were characterized and economics of sampling plans were evaluated by comparing sampling costs between spatial and conventional (non-spatial) sampling plans. Semivariogram modelling and spatial by with distance indices showed that corn rootworm adults were significantly (P < 0.05) aggregated at peak population densities and any two samples were spatially correlated within approximately 45 m, with 39–90% of the variability explained by spatial dependence. Sampling costs for spatial sampling plans linearly increased as the sampling distance decreased and exponentially increased as the field size increased. Although sampling costs for non-spatial sampling plans were generally lower, spatial sampling plans could be more economical when the mean insect density became lower and the field size became smaller. This study demonstrated that spatial sampling plans could be optimized to minimize the sampling costs and maximize the spatial resolution.     

Selection of entomopathogenic fungi for the control of the western corn rootworm Diabrotica virgifera virgifera

Abstract:  The western corn rootworm Diabrotica virgifera virgifera Le Conte (Col., Chrysomelidae), a serious pest of maize, has been recently introduced into Europe. Several approaches for its control are presently under investigation including microbial agents. During a field survey in Hungary in 2005, naturally occurring entomopathogenic fungi were found to attack this pest. These novel isolates together with standard isolates were tested for virulence against D. v. virgifera larvae and adults. Twenty strains of Metarhizium anisopliae , Beauveria bassiana and Beauveria brongniartii were used in bioassays in the laboratory. Larvae and adults were dipped into a spore suspension with a concentration of 1 × 10⁷ conidia (con.)/ml. They were kept for 14 days at 22°C (±2°C) and 70% relative humidity. The number of infected larvae and adults were counted and infection rates were calculated. Adults were significantly more susceptible to entomopathogenic fungi than larvae. The most virulent isolate infected about 47% of larvae ( M. anisopliae Ma2277), whereas the infection rate in adults was up to 97% ( M. anisopliae Ma2275). Isolates of M. anisopliae caused significantly higher mortalities than isolates of B. brongniartii and B. bassiana . Most of the adult beetles were killed within 12 days. Isolates from D. v. virgifera were more virulent than those from other hosts.     

Comparative susceptibility of western corn rootworm (Coleoptera: Chrysomelidae) adults to selected insecticides in Kansas

Susceptibility of adult populations of the western corn rootworm, Diabrotica virgifera virgifera LeConte, to several insecticides was evaluated in seven Kansas counties, including Dickinson, Ford, Finney, Pottawatomie, Republic, Riley, and Stevens, between 1996 and 2002. All populations surveyed were highly susceptible to methyl parathion with the largest difference in susceptibility of only three-fold based on 16 complete bioassays for the populations from six counties over a 5-yr period. Noticeable decreases in carbaryl susceptibility were found in populations collected from Republic County between 1997 and 2001 when the cucurbitacin-carbaryl-based bait SLAM was widely used as an areawide management approach for adult corn rootworm control. However, the lowered carbaryl susceptibility returned to previous levels 1 yr after the use of SLAM was halted in the managed (treated) cornfields. This change implies possible dispersal of insects into the relatively small managed area from surrounding untreated cornfields and / or some fitness cost associated with carbaryl resistance within the population. Relative susceptibility of western corn rootworm adults also was evaluated for seven commonly used insecticides, including bifenthrin, carbaryl, chlorpyrifos, cypermethrin, fipronil, malathion, and methyl parathion. They were tested with corn rootworm adults collected from a single cornfield. Methyl parathion and bifenthrin were highly toxic to corn rootworm adults, and cypermethrin, chlorpyrifos, carbaryl, and malathion were only slightly less toxic. Although fipronil was highly toxic to adult rootworms, its activity was much slower than that of other insecticides. Thus, bifenthrin and methyl parathion were among the most effective in killing corn rootworm adults.     

Assessment of establishment and persistence of entomopathogenic nematodes for biological control of western corn rootworm

Abstract:  The use of entomopathogenic nematodes (EPN) is potentially one ecological approach to control the invasive alien western corn rootworm ( Diabrotica virgifera virgifera LeConte, Col., Chrysomelidae) in Europe. This study investigated the establishment and the short- and long-term persistence of Heterorhabditis bacteriophora Poinar (Rhabditida: Heterorhabditidae), Heterorhabditis megidis Poinar, Jackson and Klein (Rh., Heterorhabditidae) and Steinernema feltiae Filipjev (Rh., Steinernematidae) in three maize fields in southern Hungary, using the insect-baiting technique. All three EPN species equally established and persisted in maize fields. The timing of application (April or June) did not influence the establishment of EPN species. EPNs persisted for 2–5 months, i.e. they survived up to and throughout D. v. virgifera larval occurrence in the soil. Results demonstrate that D. v. virgifera larvae can potentially be controlled by EPNs during the same year of EPN application but no long-term control effect is expected under intensive maize cultivation practices.     

Validation of a nested error component model to estimate damage caused by corn rootworm larvae

The corn rootworm complex (Coleoptera: Chrysomelidae) constitutes a significant threat to maize production in the United States, and more recently, in Europe. We conducted an analysis of readily available field trial data to validate an existing damage function for corn rootworm larvae. We used a nested error component model with unbalanced panel data to describe the relationship between yield loss and root injury caused by these insects. These data were collected by personnel with the Insect Management and Insecticide Evaluation Programme (Department of Crop Sciences, University of Illinois) and represent 19 location‐years. To our knowledge, this is the largest data set used to estimate a damage function for corn rootworm larvae. Unlike many experiments examining the relationship between root injury and yield loss caused by corn rootworm larvae, the data set used for our analysis includes many Bt maize hybrids. Our model suggests that for each node of roots injured by corn rootworm larvae, a yield loss of approximately 15% can be expected. Statistically significant variance components included an effect of location and experimental error. We speculate that variation in weather across experimental sites was the principal factor contributing to the significant effect of location. The substantial experimental error observed for our model highlights the limitations of utilizing a multi‐year, geographically diverse damage function for predicting yield loss because of root injury on a small scale. We discuss major factors contributing to the variance components estimated by our model and suggest techniques for improving future analyses of the damage function for corn rootworm larvae.     

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).     

Field screening maize germplasm for resistance and tolerance to western corn rootworms (Col.: Chrysomelidae)†

Abstract: In the hopes of lessening the current reliance on soil insecticides, developing a viable alternative for transgenic maize hybrids, and providing sustainable options for Europe, researchers recently have been developing novel maize lines that exhibit resistance and/or tolerance to corn rootworm larvae. Here we report the results of a 2‐year field experiment in a northern growing region assessing the resistance and tolerance of 10 experimental synthetic maize populations selected for varying levels of damage from western corn rootworm larvae, Diabrotica virgifera virgifera LeConte (Col.: Chrysomelidae) and four maize hybrids. Maize non‐preference, antibiosis and tolerance to rootworms was evaluated using previously established methods, including: the Iowa 1–6 root damage rating scale, root fresh weight, compensatory root growth ratings and adult rootworm emergence. Among the experimental synthetic maize populations, BS29‐11‐01 was the most susceptible, and had a mean root damage rating that was greater than the highly susceptible maize hybrid B37 × H84. This line also had the lowest mean root fresh weight and one of the lowest mean compensatory root growth ratings. In contrast, CRW8‐3 appeared to be tolerant to western corn rootworms, and had the lowest mean root damage rating, which was comparable with that of the non‐transgenic hybrid DeKalb^® 46‐26.     

Post-establishment movement of western corn rootworm larvae (Coleoptera: Chrysomelidae) in Central Missouri corn

If registered, transgenic corn, Zea mays L., with corn rootworm resistance will offer a viable alternative to insecticides for managing Diabrotica spp. corn rootworms. Resistance management to maintain susceptibility is in the interest of growers, the Environmental Protection Agency, and industry, but little is known about many aspects of corn rootworm biology required for an effective resistance management program. The extent of larval movement by the western corn rootworm, Diabrotica virgifera virgifera LeConte, that occurs from plant-to-plant or row-to-row after initial establishment was evaluated in 1998 and 1999 in a Central Missouri cornfield. Post-establishment movement by western corn rootworm larvae was clearly documented in two of four treatment combinations in 1999 where larvae moved up to three plants down the row and across a 0.46-m row. Larvae did not significantly cross a 0.91-m row after initial host establishment in 1998 or 1999, whether or not the soil had been compacted by a tractor and planter. In the current experiment, western corn rootworm larvae moved from highly damaged, infested plants to nearby plants with little to no previous root damage. Our data do not provide significant insight into how larvae might disperse after initial establishment when all plants in an area are heavily damaged or when only moderate damage occurs on an infested plant. A similar situation might also occur if a seed mixture of transgenic and isoline plants were used and if transgenic plants with rootworm resistance are not repellent to corn rootworm larvae.     

In-field labeling of western corn rootworm adults (Coleoptera: Chrysomelidae) with rubidium

Field and laboratory studies were conducted in 2000 and 2001 to determine the feasibility of mass marking western corn rootworm adults, Diabrotica virgifera virgifera LeConte, with RbCl in the field. Results showed that application of rubidium (Rb) in solution to both the soil (1 g Rb/plant) and whorl (1 g Rb/plant) of corn plants was optimal for labeling western corn rootworm adults during larval development. Development of larvae on Rb-enriched corn with this technique did not significantly influence adult dry weight or survival. Rb was also highly mobile in the plant. Application of Rb to both the soil and the whorl resulted in median Rb concentrations in the roots (5,860 ppm) that were 150-fold greater than concentrations in untreated roots (38 ppm) 5 wk after treatment. Additionally, at least 90% of the beetles that emerged during the first 3 wk were labeled above the baseline Rb concentration (5 ppm dry weight) determined from untreated beetles. Because emergence was 72% complete at this time, a significant proportion of the population had been labeled. Results from laboratory experiments showed that labeled beetles remained distinguishable from unlabeled beetles for up to 4 d postemergence. The ability to efficiently label large numbers of beetles under field conditions and for a defined period with virtually no disruption of the population provides an unparalleled opportunity to conduct mark-recapture experiments for quantifying the short-range, intrafield movement of adult corn rootworms.     

International cooperation on western corn rootworm ecology research: state-of-the-art and future research

1 Invasive pest species are challenging partly because the invasion process may be highly dynamic and because of the lack of knowledge of many researchers, professionals and farmers in the newly-invaded regions. The chrysomelid Diabrotica virgifera virgifera LeConte is such an invasive pest. It has been the main pest of continuous maize in the U.S.A. for more than 60 years and is currently spreading throughout Europe.
2 In the area with a long history of this pest (Central and North America), scientific knowledge concerning the ecology of this pest has accumulated over the last decades. This resource is of great importance to both America and Europe and has to be gathered, shared and adapted to new situations. We therefore examined, both qualitatively and quantitatively, the scientific literature relating to D.   virgifera virgifera ecology.
3 The quantitative analysis suggests that research on D.   virgifera virgifera ecology is still in its infancy in Europe and suffers from geographical barriers (between Europe and North America and between linguistic areas within Europe) and that scientific communication should be strengthened both between North America and Europe and within Europe.
4 As a first solution to this problem, we introduce three companion review articles that constitute a landmark for D.   virgifera virgifera research, enabling European and American scientists and decision-makers to orient themselves and discover new opportunities for research. We also stress that international research cooperation is the most important key to successfully manage invasive species.     

Identification of feeding stimulants in corn roots for western corn rootworm (Coleoptera: Chrysomelidae) larvae

Using a bioassay-driven approach, we have isolated and identified a blend of compounds from the roots of germinating corn, Zea mays L., that serve as feeding stimulants for neonate western corn rootworm larvae, Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae). The active blend is a combination of simple sugars (30:4:4 mg/ml glucose:fructose:sucrose in the corn root) plus at least one of the free fatty acids in germinating corn roots (2:5 mg/ml oleic acid:linoleic acid in the corn root). When an extract of germinating corn was partitioned into an ethyl acetate fraction and an aqueous fraction, full feeding occurred only when the two fractions were recombined, indicating that the phagostimulant was comprised of both polar and nonpolar components. Gas chromatography-mass spectrometry analysis of root extracts from germinating corn seedlings revealed a blend of 20 compounds from a variety of chemical classes, including small sugars, diacids, amino acids, inorganic compounds, and free fatty acids. When the major components were tested in feeding bioassays, the sugars and lipids were shown to be essential for feeding by larvae, but the two classes of compounds were only effective when combined. The sugars alone elicited feeding by only 40% of larvae, but the percentage of larvae feeding was increased significantly with the addition of linoleic acid (91.7% larvae feeding) or oleic acid (85.8% larvae feeding). The amino acids alone were not essential elements for feeding by western corn rootworm larvae.     

Inheritance of methyl-parathion resistance in Nebraska western corn rootworm populations (Coleoptera: Chrysomelidae)

Field populations of western corn rootworm, Diabrotica virgifera virgifera LeConte, were collected from three different sites (York Co., Phelps Co., and Saunders Co.) in Nebraska during 1996. Adult bioassays of these three populations were conducted with different concentrations of methyl-parathion and at a diagnostic concentration (1.0 microg/ml) to determine resistance levels among these populations. Self and reciprocal crosses were made between the two resistant and one susceptible laboratory-reared populations. Dose-responses and dominance ratios calculated for the four reciprocal crosses indicated that resistance was incompletely dominant in both strains, although in one of the strains there was an indication of sex linkage. However, evaluation of native polyacrylamide gels stained for nonspecific esterases and nonspecific esterase activity of parents and F1 progeny of the crosses suggested that esterase inheritance was completely dominant and autosomal. The results of this study were inconclusive with regard to the precise nature of inheritance, because the bioassays and esterase assays could not discriminate between heterozygotes and homozygotes. However, they do provide insight into the potential for developing simple diagnostic assays to assess resistance frequencies. Based on the inheritance studies described in this investigation, we can begin to generate information on specific genetic factors that dictate the evolutionary divergence of discrete resistant populations and facilitate modeling efforts designed to approximate the movement of genes for resistance among populations.     

Insecticide enhancement with feeding stimulants in corn for western corn rootworm larvae (Coleoptera: Chrysomelidae)

Amounts of the insecticide thiamethoxam required for 50% mortality of western corn rootworm larvae, Diabrotica virgifera virgifera LeConte, were reduced 100-fold when extracts of germinating corn, Zea mays L., were used to entice neonate larvae to feed on it. In behavioral bioassays, neonate rootworm larvae fed vigorously on filter paper disks treated with liquid pressed from corn roots. Moreover, disks treated with an acetone extract of corn (dried and rewetted with water) also elicited strong feeding from larvae. Larvae wandered away from filter paper disks treated with distilled water without feeding. Dilutions of thiamethoxam were tested in the bioassay alone or with corn extract and the efficacy of this insecticide was improved by the addition of the corn extract. For solutions containing 10 ppm thiamethoxam, 95% larval mortality occurred after 30 min of exposure when corn extract was present, but only 38% mortality occurred when the same concentration of insecticide alone (no feeding stimulants) was tested. Larval mortality after 24 h was significantly higher for corn extract-treated disks with 0.01, 0.1, 1, or 10 ppm insecticide than for the same concentrations without corn extract. Thiamethoxam did not deter larval feeding on corn extract, even at the highest concentration of thiamethoxam tested.     

Antixenosis in maize reduces feeding by western corn rootworm larvae (Coleoptera: Chrysomelidae)

SUM2162 is the first known example of a naturally occurring maize, Zea mays L., genotype with antixenosis (nonpreference) resistance to western corn rootworm, Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae), larval feeding. Behavioral responses of neonate western corn rootworm larvae were evaluated in laboratory bioassays with seven maize genotypes selected for native resistance to rootworm feeding damage. Two susceptible maize genotypes and one transgenic (Bacillus thuringiensis) maize genotype were included as controls. In soil bioassays with cut roots, no larvae entered the roots of the resistant variety SUM2162, but at least 75% of the larvae entered the roots of every other maize type. Larvae made significantly fewer feeding holes in the roots of SUM2162 than in all the other maize genotypes, except the isoline control. In feeding bioassays, larval feeding varied significantly among maize genotypes, but there was no significant difference between the resistant varieties and the susceptible controls. There were no significant differences among any of the genotypes in host recognition (search) behavior of larvae after exposure to the roots. Little variation in feeding stimulant blends was observed among maize genotypes, indicating minimal contribution to the observed antixenosis.     

Interaction between western corn rootworm (Coleoptera: Chrysomelidae) larvae and root-infecting Fusarium verticillioides

A greenhouse experiment was conducted to evaluate the effect of soil-dwelling larvae of the western corn rootworm, Diabrotica virgifera virgifera LeConte, on infection of maize roots by the mycotoxin-producing plant-pathogenic fungus, Fusarium verticillioides (Saccardo) Nirenberg (synonym=Fusarium moniliforme Sheldon). The time and order of application of F. verticillioides and western corn rootworm were varied in three different treatments to investigate the influence of timing on root colonization of F. verticillioides and western corn rootworm larval development. Root feeding by western corn rootworm larvae increased root colonization by F. verticillioides (as determined by real-time polymerase chain reaction) up to 50-fold when a high inoculum (10(7) spores/plant) of F. verticillioides was applied before western corn rootworm eggs were added. This effect was stronger the earlier F. verticillioides was applied relative to the time of western corn rootworm egg application but was only significant for the high F. verticillioides inoculum density treatment; F. verticillioides colonization was not increased when a low F. verticillioides inoculum density (10(6) spores/plant) was applied. F. verticillioides slightly suppressed larval development in that the ratio of second- to third-instar larvae was higher in treatments with F. verticillioides than without F. verticillioides. F. verticillioides reduced western corn rootworm head capsule width when applied before or simultaneously with western corn rootworm. The results of this study are discussed focusing on conditions that favor root colonization by F. verticillioides and its influence on western corn rootworm larval development.     

Monitoring western corn rootworm (Coleoptera: Chrysomelidae) susceptibility to carbaryl and cucurbitacin baits in the areawide management pilot program

Areawide pest management involves the uniform application of a pest control strategy over wide geographic areas. Therefore, these programs are likely to impose intense selective pressures, and the risk for resistance development among pest species for which areawide management programs are implemented is likely to be high. Pilot studies for areawide management of western corn rootworm, Diabrotica virgifera virgifera LeConte, were conducted from 1996 to 2002 at four different sites across the Corn Belt. This program used cucurbitacin baits to deliver high doses of a traditional neurotoxic insecticide (carbaryl) to individual insects while reducing the overall rate of insecticide use. Because of the concern and potential for resistance evolution, annual assessments of susceptibility to the active ingredient carbaryl were conducted both within the managed area as well as from untreated control areas. Significantly reduced susceptibility to carbaryl based on survival at a diagnostic concentration was detected in three of the four management sites (Kansas, Iowa, and Illinois/Indiana), whereas susceptibility of beetles collected outside the managed areas remained unchanged. Additionally, significantly reduced responsiveness to cucurbitacin baits was observed in beetles collected from the managed area relative to the control area at the same three sites. These results suggest strongly that areawide management has the potential to select for resistance and that a strategy for managing resistance and reducing selective pressure should be proactively implemented.     

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.     

Assessing risk of resistance to aerial applications of methyl-parathion in western corn rootworm (Coleoptera: Chrysomelidae)

We validated a stochastic model of the evolution of resistance to adulticidal sprays of methyl-parathion in western corn rootworm, Diabrotica virgifera virgifera LeConte, populations in Nebraska. The population dynamics predicted by the model resembled that reported for field populations, and time until control failures occurred closely matched reports by commercial crop consultants. We incorporated uncertainty about the values used for 18 model parameters by replacing default values with random draws taken from a normal distribution. One parameter, the initial resistance allele frequency, was no longer measurable because of the evolution of resistance. We therefore proposed five candidate initial allele frequencies and developed probability distributions for the time to resistance for each by running 1000 simulations with parameters randomly varied. These distributions included variation because of stochastic effects as well as parameter uncertainty. We used Bayesian inference to estimate the candidate frequency most likely, given reported times to field control failures. The initial allele frequency of 10(-4) was most likely (29%), 10(-3) was less likely (28%), whereas 10(-6) was relatively unlikely (5%). Results from sensitivity analysis depended upon how evolution of resistance was measured. When resistance was examined as a genetic phenomenon, the rate of increase of the resistance allele depended almost entirely on genetic factors (LC50 values), the characteristics of the pesticide (residual activity), and the variance associated with emergence of adults. When resistance was measured as failure of methyl-parathion to reduce populations below threshold levels (0.5 gravid females per plant), parameters that contributed to population growth rate (mortality and fecundity) were also important. These data suggest two important phases in resistance evolution in corn rootworms: a genetic phase associated with negative growth rates and rapid changes in resistance allele frequencies and a rebound phase associated with positive growth rates and near fixation of the resistance allele.