Field-Evolved Resistance of the Fall Armyworm (Lepidoptera: Noctuidae) to Synthetic Insecticides in Puerto Rico and Mexico.

The fall armyworm, Spodoptera frugiperda (J. E. Smith), is one of the main pests of corn in many areas of the American continent. The reliance on pesticides to control fall armyworm has led to the development of insecticide resistance in many regions. We determined the resistance levels of fall armyworm to insecticides of different modes of action in fall armyworm populations from Puerto Rico and several Mexican states with different insecticide use patterns. Mexican populations that expressed higher resistance ratios (RR50) were: Sonora (20-fold to chlorpyriphos), Oaxaca (19-fold to permethrin), and Sinaloa (10-fold to flubendamide). The Puerto Rico population exhibited a remarkable field-evolved resistance to many pesticides. The RR50 to the insecticides tested were: flubendiamide (500-fold), chlorantraniliprole (160-fold), methomyl (223-fold), thiodicarb (124-fold), permethrin (48-fold), chlorpyriphos (47-fold), zeta-cypermethrin (35-fold), deltamethrin (25-fold), triflumuron (20-fold), spinetoram (14-fold). Spinosad (eightfold), emamectin benzoate and abamectin (sevenfold) displayed lower resistance ratio. However, these compounds are still effective to manage fall armyworm resistance in Puerto Rico. Fall armyworm populations from Mexico show different levels of susceptibility, which may reflect the heterogeneity of the pest control patterns in this country. The status of insecticide resistance in the fall armyworm from Puerto Rico indicates a challenging situation for the control of this pest with these insecticides in the close future. Lessons learned from this research might be applied in regions with recent invasions of fall armyworm in Africa.

The Red Queen in a potato field: integrated pest management versus chemical dependency in Colorado potato beetle control.

Originally designed to reconcile insecticide applications with biological control, the concept of integrated pest management (IPM) developed into the systems-based judicious and coordinated use of multiple control techniques aimed at reducing pest damage to economically tolerable levels. Chemical control, with scheduled treatments, was the starting point for most management systems in the 1950s. Although chemical control is philosophically compatible with IPM practices as a whole, reduction in pesticide use has been historically one of the main goals of IPM practitioners. In the absence of IPM, excessive reliance on pesticides has led to repeated control failures due to the evolution of resistance by pest populations. This creates the need for constant replacement of failed chemicals with new compounds, known as the 'insecticide treadmill'. In evolutionary biology, a similar phenomenon is known as the Red Queen principle - continuing change is needed for a population to persevere because its competitors undergo constant evolutionary adaptation. The Colorado potato beetle, Leptinotarsa decemlineata (Say), is an insect defoliator of potatoes that is notorious for its ability to develop insecticide resistance. In the present article, a review is given of four case studies from across the United States to demonstrate the importance of using IPM for sustainable management of a highly adaptable insect pest. Excessive reliance on often indiscriminate insecticide applications and inadequate use of alternative control methods, such as crop rotation, appear to expedite evolution of insecticide resistance in its populations. Resistance to IPM would involve synchronized adaptations to multiple unfavorable factors, requiring statistically unlikely genetic changes. Therefore, integrating different techniques is likely to reduce the need for constant replacement of failed chemicals with new ones.

Defining terms for proactive management of resistance to Bt crops and pesticides.

Evolution of pest resistance to pesticides is an urgent global problem with resistance recorded in at least 954 species of pests, including 546 arthropods, 218 weeds, and 190 plant pathogens. To facilitate understanding and management of resistance, we provide definitions of 50 key terms related to resistance. We confirm the broad, long-standing definition of resistance, which is a genetically based decrease in susceptibility to a pesticide, and the definition of "field-evolved resistance," which is a genetically based decrease in susceptibility to a pesticide in a population caused by exposure to the pesticide in the field. The impact of field-evolved resistance on pest control can vary from none to severe. We define "practical resistance" as field-evolved resistance that reduces pesticide efficacy and has practical consequences for pest control. Recognizing that resistance is not "all or none" and that intermediate levels of resistance can have a continuum of effects on pest control, we describe five categories of field-evolved resistance and use them to classify 13 cases of field-evolved resistance to five Bacillus thuringiensis (Bt) toxins in transgenic corn and cotton based on monitoring data from five continents for nine major pest species. We urge researchers to publish and analyze their resistance monitoring data in conjunction with data on management practices to accelerate progress in determining which actions will be most useful in response to specific data on the magnitude, distribution, and impact of resistance.

Geographic variation in cannibalism in Colorado potato beetle (Coleoptera: Chrysomelidae) populations.

Cannibalism can have a large effect on population growth and survival in stressful environments, possibly including those created by insecticide use. In this study, we collected Colorado potato beetles from three isolated areas in the northeastern United States known for high levels of resistance to neonicotinoids. We measured resistance to imidacloprid in each of those populations, a laboratory susceptible population, and in hybrids between the three field populations and the laboratory susceptible population. We fed neonates eggs from resistant dams fed either imidacloprid-treated or untreated foliage to determine whether cannibals are exposed to toxins sequestered in eggs. We measured egg cannibalism by hatchlings within the clutch in each population and hybrids, and examined how fecundity and several variables associated with egg development varied among populations and with cannibalism, to see which traits might enhance or reduce cannibalism. Cannibalism varied significantly among populations, accounting for most of the variation in hatching success. Variability in egg development time and hatch rate in the absence of cannibalism in some populations affected rates of cannibalism. Resistance varied significantly among the field populations but was not related to cannibalism. Neonates fed eggs from dams on treated foliage showed signs of intoxication or death. Cannibalism appears to be part of a varying life history strategy in this species, with some populations laying larger and more cannibalistic clutches and the New York population laying smaller clutches with higher hatching success owing to reduced cannibalism.

Dose-response relationships for the antifeedant effects of Humulus lupulus extracts against larvae and adults of the Colorado potato beetle.

BACKGROUND: Dose-response relationships for antifeedant effects of H. lupulus extracts against larvae and adults of the Colorado potato beetle (CPB) were determined in laboratory conditions. The larval and adult beetles were fed on potato leaflets treated with H. lupulus extract ranging from 0.4 to 40 mg mL(-1) in a no-choice situation. Their feeding behavior was recorded, and larval growth and antifeedant indexes were calculated. RESULTS: H. lupulus treatments significantly affected larval growth rate, and at higher concentrations the larval weights were significantly reduced over the course of the assay. Adults of CPB were more sensitive to the extracts than the larvae, and, even at lower doses, adult beetles were arrested for longer periods than larvae. CONCLUSION: These results indicate that H. lupulus extracts may have potential for control of CPB, particularly in organic farms where conventional insecticides are not available.

Novaluron causes reduced egg hatch after treating adult codling moths, Cydia pomenella: support for transovarial transfer.

The codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), is a primary pest of apples throughout the United States. Reliance on broad spectrum organophosphates has been declining with the slated cancellation and has shifted towards narrow spectrum insecticides. Novaluron, a chitin synthesis inhibitor, has primarily been used for its ovicidal and larvacidal activities. However, recent studies have demonstrated a transovarial effect after exposure to adults. The effects of novaluron were studied to determine if reduced egg hatch occurs after exposure of different sexes to this compound. Effects of this compound through horizontal transfer were also compared with a topical application to C. pomonella eggs. Results from independent exposure of different sexes to novaluron were different than the control for all three exposure types; male only, female only, and both treated. The horizontal transfer experiment yielded no significant difference while the topical application of novaluron on eggs showed significantly lower egg hatch. Although novaluron has no direct toxicity to adults, the results of this study demonstrate that the delayed lethal activity of this compound reduces hatching of eggs laid by treated adults. Along with the direct ovicidal and larvicidal properties of novaluron, the delayed lethal activity provides an important contribution to the overall control seen in the field.

Modeling protected species habitat and assigning risk to inform regulatory decisions.

In the United States, environmental regulatory agencies are required to use "best available" scientific information when making decisions on a variety of issues. However, agencies are often hindered by coarse or incomplete data, particularly as it pertains to threatened and endangered species protection. Stakeholders often agree that more resolute and integrated processes for decision-making are desirable. We demonstrate a process that uses species occurrence data for a federally endangered insect (Karner blue butterfly), a readily available habitat modeling tool, and spatially explicit information about an important Michigan commodity (tart cherries). This case study has characteristics of many protected species regulatory decisions in that species occurrence data were sparse and unequally distributed; regulatory decisions (on pesticide use) were required with potentially significant impacts on a viable agricultural industry; and stakeholder relations were diverse, misinformed, and, in some situations, unjustly contentious. Results from our process include a large-scale, empirically derived habitat suitability map for the focal species and a risk ranking of tart cherry orchards with risk based on the likelihood that pesticide applications will influence the focal protected species. Although the majority (77%) of pesticide-influence zones overlapped Karner blue butterfly habitat, risk scores associated with each orchard were low. Through our process we demonstrated that spatially explicit models can help stakeholders visualize and quantify potential protected species effects. In addition, model outputs can serve to guide field activities (e.g., species surveys and implementation of pesticide buffer zones) that help minimize future effects.

Reduced egg viability in codling moth Cydia pomonella (L.) (Lepidoptera: Tortricidae) following adult exposure to novaluron.

BACKGROUND: The codling moth, Cydia pomonella (L.), is one of the principal pests of pome fruits in the world. The effects of novaluron, a benzoylurea chitin synthesis inhibitor insecticide registered for use on apples in the USA, on fecundity and egg viability in codling moth were studied under laboratory conditions. Three different exposure methods were investigated: ingestion, contact and topical spray. Additionally, the duration of novaluron sublethal effects was measured subsequent to the three modes of exposure. RESULTS: The fecundity of codling moth adults was not significantly affected by novaluron with any of the exposure methods. However, novaluron did cause significant reductions in the proportion of egg hatch with all three exposure methods. The duration of sublethal effects was 9 days or more for all modes of exposure, but with the topical spray these effects began to diminish after 6 days. CONCLUSION: Novaluron does not affect fecundity in codling moth, but has significant sublethal activity by reducing egg viability subsequent to adult exposure. The topical, contact and ingestion exposures all induce sublethal effects after exposure, and these persist to various degrees throughout codling moth oviposition. A more complete understanding of novaluron's lethal and sublethal activities will help IPM practitioners optimize its use for management of the codling moth.

Novel life stage targets against plum curculio, Conotrachelus nenuphar (Herbst), in apple integrated pest management.

In this study the authors employed the plant-insect-chemistry (PIC) triad to investigate two novel life stage targets against the plum curculio (PC), Conotrachelus nenuphar (Herbst), in apple integrated pest management (IPM). Laboratory treated apple bioassays were used to determine if the insect growth regulator (IGR) insecticides novaluron and tebufenozide have physiological effects on PC larvae following adult exposure. Curative activity bioassays were conducted for IGR, neonicotinoid, oxidiazine and organophosphate insecticides on PC larvae post-infestation, and fruit penetration profiles of insecticides were developed. The results revealed that novaluron exhibits activity on PC larvae via vertical transmission following exposure of mated females to treated substrate. Surface treatments of azinphos-methyl, thiacloprid and thiamethoxam to preinfested fruit resulted in significant reductions in larval emergence. For all compounds, 50% or more of the total recovered active ingredient was from apple skin, and for azinphos-methyl, indoxacarb and thiamethoxam it was greater than 80%. For azinphos-methyl, novaluron, methoxyfenozide and thiacloprid, however, active ingredient was recovered from every section of the apple, from skin to core. Implications for twenty-first century IPM are discussed.

Pyriproxyfen activates reproduction in prediapause northern strain plum curculio (Conotrachelus nenuphar Herbst).

Field-collected, prediapause northern strain plum curculio, Conotrachelus nenuphar (Herbst), adults were treated with the juvenile hormone analogue pyriproxyfen to assess effects on reproductive development. Adults of this pest have an obligate winter reproductive diapause and do not reach reproductive maturity until after spring emergence. Topical (1.0 microL) doses of 10, 1.0 and 0.1 microg microL(-1) pyriproxyfen induced oocyte development and reproductive maturation in all treated females as assessed by dissection. There was no increased mortality in treated beetles, and control insects showed no reproductive maturation. Treatment of prediapause males and females with 1.0 microg microL(-1) of topical pyriproxyfen or exposure to residues on fruit induced successful egg laying and F1 emergence; F1 pupation success in topical treatments and residue exposure was 47 and 59% respectively. Filial adults require re-exposure to initiate reproductive development. Treatment protocols with pyriproxyfen will allow researchers to culture the northern strain, instead of relying exclusively on the non-diapausing southern strain.

Aggregation pheromone of Metamasius spinolae (Coleoptera: Curculionidae): chemical analysis and field test.

Male Metamasius spinolae (Gylh.) produce several volatile compounds that are likely constituents of its aggregation pheromone. These compounds were identified by volatile collections and gas chromatography (GC), followed by coupled gas chromatography-mass spectrometry (GC-MS), as 2-methyl-4-heptanone [1], 6-methyl-2hepten-4-one [2], and 2-hydroxy-2-methyl-4-heptanone [3]. Preliminary field experiments using synthetic racemates of these compounds showed that significantly more adult cactus weevils were caught in traps baited with the major single compound three or the 2 + 3 binary combination than in unbaited control traps. However, highest trap efficacy occurred with the 1 + 2 binary combination and a blend of all three synthetic compounds plus prickly pear. Potential uses for the cactus weevil pheromone and possible ways to increase trap captures are discussed.

Behavioural response of Colorado potato beetle (Leptinotarsa decemlineata) larvae to selected plant extracts.

Potato leaves were treated with 2, 20 or 200 g kg(-1) solutions of extracts of five plant species collected in Turkey, and then exposed to larvae of Colorado potato beetle, Leptinotarsa decemlineata (Say). During the first 24 h of exposure, leaf consumption was not affected by 2 g kg(-1) extracts, whereas significantly more leaf tissue remained on leaves treated with 20 g kg(-1) extracts of Arctium lappa L., Bifora radians M Bieb, Humulus lupulus L. or Xanthium strumarium L. than on untreated control leaves. Feeding was not affected by the 20 g kg(-1) extract of Verbascum songaricum Schrenk ex Fisch & Mey. Extracts of all species at 200 g kg(-1) reduced larval feeding, with H. lupulus and X. strumarium providing the greatest protection. Observations of larval behaviour over the first 15 min of exposure to these extracts revealed that the interaction of beetles with leaf tissue was significantly affected by plant extracts. Feeding frequency was not affected by 2 g kg(-1) extracts but was reduced by all higher concentrations. Feeding was inhibited completely by 20 g kg(-1) of H. lupulus extract and reduced significantly compared with the controls by all other extracts. Suppression of feeding was caused by all extracts at 200 g kg(-1), with V. songaricum providing 91% reduction in feeding duration. Rejection behaviour, in which larvae did not return to the leaf after their interaction with it, was rare on 2 g kg(-1) extracts but seen in over 60% of larvae on 20 g kg(-1) extracts and over 80% on 200 g kg(-1) extracts. The present results demonstrate that these extracts have significant ability to protect potato leaves for up to 24 h by prevention of feeding behaviour by L. decemlineata. Further studies are needed to determine the potential of these plant extracts, or their active components, for use in biologically based pest management strategies.

Use of residue profile analysis to identify modes of insecticide activity contributing to control of plum curculio in apples.

Residue profile analysis techniques were developed, along with laboratory and field-based bioassays to describe the modes of insecticidal activity responsible for the control of the plum curculio, Conotrachelus nenuphar (Herbst), in apples (Malus spp.). Adult plum curculios were treated in laboratory topical bioassays to determine acute contact activity and lethal time for five insecticides. Azinphosmethyl had the highest levels of toxicity and shortest lethal time values, followed by the neonicotinoids thiacloprid, thiamethoxam, and imidacloprid, whereas indoxacarb had the highest LD50 and LT50 values for topical exposure. Field-based residual activity bioassays assessed adult mortality, and fruit and leaf injury from plum curculio exposed to 4 h, 7 d, and 14 d field-aged residues. All compounds caused significant levels of mortality to plum curculio when adults were exposed to fruit clusters 4 h post-application. Thiacloprid, thiamethoxam, and imidacloprid showed oviposition deterrence, antifeedant, and repellency effects in the 7- and/or 14 d residual bioassays and protected fruit in the absence of significant lethal activity. Indoxacarb maintained lethal activity throughout the study intervals, with the incidence of plum curculio feeding, suggesting that ingestion is an important mode of entry. For the neonicotinoids thiacloprid, thiamethoxam and imidacloprid plum curculio mortality was highly correlated with fruit and leaf surface residues. As surface residues declined, sublethal effects such as oviposition deterrence and antifeedant effect remained. The value of the plant-insect-chemistry triad model for describing the temporal dimensions of insecticidal modes of activity and understanding a compound's critical performance characteristics is discussed.

Reproductive development of northern and southern strains of plum curculio (Coleoptera: Curculionidae).

Laboratory-reared southern and field-collected northern strains of plum curculio, Conotrachelles nenuphar (Herbst), were sampled to examine the relationship between degree-day (DD) accumulation and female reproductive development, as measured by mating status, oocyte size, and number of oocytes. The overall goal was to generate an objective degree-day model for predicting damage potential that could be applied to various host commodities rather than relying on separate biofix models for each crop. Adult beetles were dissected to measure mating status, maximum oocyte size, and number of oocytes. Southern strain beetles reared at 25 degrees C initiated mating 9 d after eclosion and did not require mating to induce oocyte development. By 20 d posteclosion, unmated females had significantly higher egg loads compared with mated females of the same age. Logistic regression analysis suggests that southern and northern strain beetles had a stable maximum oocyte length of 62 and 72 microm, respectively. Northern strain females mated after overwintering; with approximately 95% of the female population mated after 134 DD (base 10 degrees C), which is before fruit set in many host crops. Oocyte size was the only measured parameter of field reproductive progress that could be linked with confidence to degree-day accumulation. The other two parameters do not share an exclusive relationship with degree-days. Rapid assessment of field-caught female reproductive status could assist in determining the potential for plum curculio damage in high-value commodities and allow for more informed control decisions.

The tomato homolog of CORONATINE-INSENSITIVE1 is required for the maternal control of seed maturation, jasmonate-signaled defense responses, and glandular trichome development.

Jasmonic acid (JA) is a fatty acid-derived signaling molecule that regulates a broad range of plant defense responses against herbivores and some microbial pathogens. Molecular genetic studies in Arabidopsis have established that JA also performs a critical role in anther and pollen development but is not essential for other developmental aspects of the plant's life cycle. Here, we describe the phenotypic and molecular characterization of a sterile mutant of tomato (jasmonic acid-insensitive1 [jai1]) that is defective in JA signaling. Although the mutant exhibited reduced pollen viability, sterility was caused by a defect in the maternal control of seed maturation, which was associated with the loss of accumulation of JA-regulated proteinase inhibitor proteins in reproductive tissues. jai1 plants exhibited several defense-related phenotypes, including the inability to express JA-responsive genes, severely compromised resistance to two-spotted spider mites, and abnormal development of glandular trichomes. We demonstrate that these defects are caused by the loss of function of the tomato homolog of CORONATINE-INSENSITIVE1 (COI1), an F-box protein that is required for JA-signaled processes in Arabidopsis. These findings indicate that the JA/COI1 signaling pathway regulates distinct developmental processes in different plants and suggest a role for JA in the promotion of glandular trichome-based defenses.

Bt: mode of action and use.

The insecticidal toxins from Bacillus thuringiensis (Bt) represent a class of biopesticides that are attractive alternatives to broad-spectrum "hard" chemistries. The U.S. Food Quality Protection Act and the European Economic Council directives aimed at reducing the use of carbamate and organophosphate insecticides were expected to increase the use of narrowly targeted, "soft" compounds like Bt. Here we summarize the unique mode of action of Bt, which contributes to pest selectivity. We also review the patterns of Bt use in general agriculture and in specific niche markets. Despite continued predictions of dramatic growth for biopesticides due to US Food Quality Protection Act-induced cancellations of older insecticides, Bt use has remained relatively constant, even in niche markets where Bt has traditionally been relatively high.