Development of a Novel Dry, Sticky Trap Design Incorporating Visual Cues for Drosophila suzukii (Diptera: Drosophilidae).

Drosophila suzukii Matsumura (Diptera: Drosophilidae) is currently one of the most serious invasive pests for berry crops and cherries worldwide. The development of an effective monitoring trap that is reliable at detecting small populations to guide management decisions is greatly needed. To develop a novel dry trapping system, D. suzukii were trapped under field conditions in cherry orchards and raspberry high tunnels using various baited dry trap designs that were compared with the currently available deli-cup style traps that utilize a liquid bait or drowning solution. In a test in cherry orchards, red panel and combination yellow panel plus red sphere traps captured significantly more flies than yellow panel traps when all were baited with a Scentry lure. In a separate test in cherry, red sphere traps with the Scentry lure captured significantly more flies than the deli-cup traps with the Scentry lure or with the yeast sugar bait, and red panel traps with the Scentry lure captured significantly more flies than deli-cup traps with the Scentry lure. In raspberry high tunnels, red sphere traps with the Scentry lure captured significantly more flies than deli-cup traps with the Scentry lure. Red traps baited with the same lure as clear deli-cup traps consistently captured more D. suzukii, demonstrating that traps integrating a visual cue in combination with an olfactory cue are superior tools for monitoring D. suzukii. A dry trap requires less labor and maintenance than cup traps containing a liquid, improving the ease of use of D. suzukii monitoring traps.

Line-Trapping of Codling Moth (Lepidoptera: Tortricidae): A Novel Approach to Improving the Precision of Capture Numbers in Traps Monitoring Pest Density.

This field study of codling moth, Cydia pomonella (L.), response to single versus multiple monitoring traps baited with codlemone demonstrates that precision of a given capture number is alarmingly poor when the population is held constant by releasing moths. Captures as low as zero and as high as 12 males per single trap are to be expected where the catch mode is three. Here, we demonstrate that the frequency of false negatives and overestimated positives for codling moth trapping can be substantially reduced by employing the tactic of line-trapping, where five traps were deployed 4 m apart along a row of apple trees. Codling moth traps spaced closely competed only slightly. Therefore, deploying five traps closely in a line is a sampling technique nearly as good as deploying five traps spaced widely. But line trapping offers a substantial savings in time and therefore cost when servicing aggregated versus distributed traps. As the science of pest management matures by mastering the ability to translate capture numbers into estimates of absolute pest density, it will be important to employ a tactic like line-trapping so as to shrink the troublesome variability associated with capture numbers in single traps that thwarts accurate decisions about if and when to spray. Line-trapping might similarly increase the reliability and utility of density estimates derived from capture numbers in monitoring traps for various pest and beneficial insects.

Genomic innovations, transcriptional plasticity and gene loss underlying the evolution and divergence of two highly polyphagous and invasive Helicoverpa pest species.

BACKGROUND: Helicoverpa armigera and Helicoverpa zea are major caterpillar pests of Old and New World agriculture, respectively. Both, particularly H. armigera, are extremely polyphagous, and H. armigera has developed resistance to many insecticides. Here we use comparative genomics, transcriptomics and resequencing to elucidate the genetic basis for their properties as pests. RESULTS: We find that, prior to their divergence about 1.5 Mya, the H. armigera/H. zea lineage had accumulated up to more than 100 more members of specific detoxification and digestion gene families and more than 100 extra gustatory receptor genes, compared to other lepidopterans with narrower host ranges. The two genomes remain very similar in gene content and order, but H. armigera is more polymorphic overall, and H. zea has lost several detoxification genes, as well as about 50 gustatory receptor genes. It also lacks certain genes and alleles conferring insecticide resistance found in H. armigera. Non-synonymous sites in the expanded gene families above are rapidly diverging, both between paralogues and between orthologues in the two species. Whole genome transcriptomic analyses of H. armigera larvae show widely divergent responses to different host plants, including responses among many of the duplicated detoxification and digestion genes. CONCLUSIONS: The extreme polyphagy of the two heliothines is associated with extensive amplification and neofunctionalisation of genes involved in host finding and use, coupled with versatile transcriptional responses on different hosts. H. armigera's invasion of the Americas in recent years means that hybridisation could generate populations that are both locally adapted and insecticide resistant.

Maximizing Information Yield From Pheromone-Baited Monitoring Traps: Estimating Plume Reach, Trapping Radius, and Absolute Density of Cydia pomonella (Lepidoptera: Tortricidae) in Michigan Apple.

Novel methods of data analysis were used to interpret codling moth (Cydia pomonella) catch data from central-trap, multiple-release experiments using a standard codlemone-baited monitoring trap in commercial apple orchards not under mating disruption. The main objectives were to determine consistency and reliability for measures of: 1) the trapping radius, composed of the trap's behaviorally effective plume reach and the maximum dispersive distance of a responder population; and 2) the proportion of the population present in the trapping area that is caught. Two moth release designs were used: 1) moth releases at regular intervals in the four cardinal directions, and 2) evenly distributed moth releases across entire approximately 18-ha orchard blocks using both high and low codling moth populations. For both release designs, at high populations, the mean proportion catch was 0.01, and for the even release of low populations, that value was approximately 0.02. Mean maximum dispersive distance for released codling moth males was approximately 260 m. Behaviorally effective plume reach for the standard codling moth trap was < 5 m, and total trapping area for a single trap was approximately 21 ha. These estimates were consistent across three growing seasons and are supported by extraordinarily high replication for this type of field experiment. Knowing the trapping area and mean proportion caught, catch number per single monitoring trap can be translated into absolute pest density using the equation: males per trapping area = catch per trapping area/proportion caught. Thus, catches of 1, 3, 10, and 30 codling moth males per trap translate to approximately 5, 14, 48, and 143 males/ha, respectively, and reflect equal densities of females, because the codling moth sex ratio is 1:1. Combined with life-table data on codling moth fecundity and mortality, along with data on crop yield per trapping area, this fundamental knowledge of how to interpret catch numbers will enable pest managers to make considerably more precise projections of damage and therefore more precise and reliable decisions on whether insecticide applications are justified. The principles and methods established here for estimating absolute codling moth density may be broadly applicable to pests generally and thereby could set a new standard for integrated pest management decisions based on trapping.

Alightment of Spotted Wing Drosophila (Diptera: Drosophilidae) on Odorless Disks Varying in Color.

Methods for trapping spotted wing drosophila, Drosophila suzukii (Matsmura) (Diptera: Drosophilidae), have not yet been optimized for detecting this devastating pest of soft-skinned fruits. Here, we report outcomes of choice and no-choice laboratory bioassays quantifying the rates of spotted wing drosophila alightment on 5-cm-diameter sticky disks of various colors, but no fruit odors. Red, purple, and black disks captured the most spotted wing drosophila when presented against a white background. Male and female spotted wing drosophila responded identically in these tests. Significantly more D. suzukii were captured on the red and yellow disks than those presenting the corresponding grayscale for that color, proving that D. suzukii perceives colors and not just the level of target brightness. Fluorescent red is the best candidate for trap color, while clear and white are the least desirable. However, when the background was switched to black, all nonfluorescent colors were equally acceptable to spotted wing drosophila, suggesting that background must be specified when reporting spotted wing drosophila color preference. Additional spotted wing drosophila research is justified on the effects of target color against natural backgrounds.

Divergent host preferences of above- and below-ground Culex pipiens mosquitoes and their hybrid offspring.

Culex pipiens form pipiens and Cx. pipiens form molestus (Diptera: Culicidae) belong to a cosmopolitan taxonomic group known as the Pipiens Assemblage. Hybridization between these forms is thought to contribute to human transmission of West Nile virus (WNV) in North America. Complementary choice and no-choice landing assays were developed to examine host acceptance by North American Cx. pipiens in the laboratory. Populations collected from above- and below-ground sites in suburban Chicago were identified as forms pipiens and molestus using a polymerase chain reaction-based assay. Avian and human host acceptance was then quantified for the two populations, as well as for their hybrid and backcross offspring. No-choice tests were used to demonstrate that both the pipiens and molestus forms were capable of feeding on human and avian hosts. Choice tests were used to demonstrate that form pipiens females were strongly avian-seeking; an individual's probability of accepting the chick host was 85%. Form molestus females were more likely to accept the human host (87%). Rates of host acceptance by F1 and backcross progeny were intermediate to those of their parents. The results suggest that host preferences in Cx. pipiens are genetically determined, and that ongoing hybridization between above- and below-ground populations is an important contributor to epizootic transmission of WNV in North America.

Application of a reverse dot blot DNA-DNA hydridization method to quantify host-feeding tendencies of two sibling species in the Anopheles gambiae complex.

A DNA-DNA hybridization method, reverse dot blot analysis (RDBA), was used to identify Anopheles gambiae s.s. and Anopheles arabiensis (Diptera: Culicidae) hosts. Of 299 blood-fed and semi-gravid An. gambiae s.l. collected from Kisian, Kenya, 244 individuals were identifiable to species; of these, 69.5% were An. arabiensis and 29.5% were An. gambiae s.s. Host identifications with RDBA were comparable with those of conventional polymerase chain reaction (PCR) followed by direct sequencing of amplicons of the vertebrate mitochondrial cytochrome b gene. Of the 174 amplicon-producing samples used to compare these two methods, 147 were identifiable by direct sequencing and 139 of these were identifiable by RDBA. Anopheles arabiensis bloodmeals were mostly (94.6%) bovine in origin, whereas An. gambiae s.s. fed upon humans more than 91.8% of the time. Tests by RDBA detected that two of 112 An. arabiensis contained blood from more than one host species, whereas PCR and direct sequencing did not. Recent use of insecticide-treated bednets in Kisian is likely to have caused the shift in the dominant vector species from An. gambiae s.s. to An. arabiensis. Reverse dot blot analysis provides an opportunity to study changes in host-feeding by members of the An. gambiae complex in response to the broadening distribution of vector control measures targeting host-selection behaviours.

General principles of attraction and competitive attraction as revealed by large-cage studies of moths responding to sex pheromone.

Knowledge of how insects are actually affected by sex pheromones deployed throughout a crop so as to disrupt mating has lacked a mechanistic framework sufficient for guiding optimization of this environmentally friendly pest-control tactic. Major hypotheses are competitive attraction, desensitization, and camouflage. Working with codling moths, Cydia pomonella, in field cages millions of times larger than laboratory test tubes and at substrate concentrations trillions of times less than those typical for enzymes, we nevertheless demonstrate that mating disruption sufficiently parallels enzyme (ligand) -substrate interactions so as to justify adoption of conceptual and analytical tools of biochemical kinetics. By doing so, we prove that commercial dispensers of codling moth pheromone first competitively attract and then deactivate males probably for the remainder of a night. No evidence was found for camouflage. We generated and now validate simple algebraic equations for attraction and competitive attraction that will guide optimization and broaden implementation of behavioral manipulations of pests. This analysis system also offers a unique approach to quantifying animal foraging behaviors and could find applications across the natural and social sciences.

Toxicity of bloodmeals from ivermectin-treated cattle to Anopheles gambiae s.l.

Two anthelmintic drugs used as cattle dewormers, ivermectin and moxidectin, were tested for their lethal and sublethal effects on the malarial vectors Anopheles gambiae s.s. and An. arabiensis. In the laboratory, direct addition of ivermectin to bovine blood reduced the survivorship and fecundity of mosquitoes fed on the blood. The median lethal concentration (LC(50)) of ivermectin in the bloodmeal, for the laboratory populations of An. gambiae s.l., was 19.8 ppb. In the field, commercially available formulations containing ivermectin or moxidectin were injected into cattle at three times the recommended dose. Most (90%) of the An. gambiae s.s. that fed on the ivermectin-treated cattle within 2 weeks of treatment failed to survive more than 10 days post-bloodmeal. No eggs were deposited by An. gambiae s.s. that fed on ivermectin-treated cattle within 10 days of treatment. In contrast, the survivorship and egg production of the mosquitoes that fed on the moxidectin-treated cattle were no different from those feeding on untreated cattle. These results indicate that treatment of cattle with ivermectin could be used, as part of an integrated control programme, to reduce the zoophilic vector populations that contribute to the transmission of the parasites responsible for human malaria.

Designation of chemicals in terms of the locomotor responses they elicit from insects: an update of Dethier et al. (1960).

A scheme updating that of Dethier et al. (1960) (J. Econ. Entomol. 53: 134-136) for chemicals influencing insect locomotor behavior is introduced. Attractant, repellent, and arrestant retain their previous definitions. However, attractants or repellents are now recognized to operate both by kinetic and tactic mechanisms. Locomotor initiator is a new term for stimuli that activate normal levels of kinetic locomotion. Locomotor stimulant is reserved for activation of abnormally high kinetic locomotion, like that arising upon sublethal exposure to certain insecticides. The new terms engagent and disengagent apply to chemicals that, by their effects on locomotion, increase or decrease interaction with the source of stimulation, respectively. With these clarifications, insect behavioral terms unique to medical entomology but contradicting Dethier et al.'s classical scheme can be reconciled with the vocabulary of formal behavioral science.

Mating disruption of citrus leafminer mediated by a noncompetitive mechanism at a remarkably low pheromone release rate.

The citrus leafminer, Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae), is a worldwide pest of citrus. A season-long investigation was conducted that evaluated mating disruption for this pest. Effective disruption of the male P. citrella orientation to pheromone traps (98%) and reduced flush infestation by larvae was achieved for 221 d with two deployments of a 3:1 blend of (Z,Z,E)-7,11,13-hexadecatrienal/(Z,Z)-7,11-hexadecadienal at a remarkably low rate of 1.5 g active ingredient (AI)/ha per deployment. To gain insight into the mechanism that mediates the disruption of P. citrella, male moth catch was quantified in replicated plots of citrus treated with varying densities of pheromone dispensers. The densities of septum dispensers compared were: 0 (0/ha, 0.0 g AI/ha), 0.2 (one every fifth tree or 35/ha, 0.05 g AI/ha), 1 (215/ha, 0.29 g AI/ha), and 5 per tree (1,100/ha, 1.5 g AI/ha). Profile analysis by previously published mathematical methods matched predictions of noncompetitive mating disruption. Behavioral observations of male P. citrella in the field revealed that males did not approach mating disruption dispensers in any of the dispenser density treatments. The current report presents the first set of profile analyses combined with direct behavioral observations consistent with previously published theoretical predictions for a noncompetitive mechanism of mating disruption. The results suggest that disruption of P. citrella should be effective even at high population densities given the density-independent nature of disruption for this species and the remarkably low rate of pheromone per hectare required for efficacy.

Alcohol consumption among recreational boaters: factors for intervention.

Recreational boating is a popular leisure time activity in many countries. It is estimated that, in Australia, boating incidents cause more harm than rail and air crashes combined and, in terms of transport, are second only to motor vehicle crashes as a cause of serious injury. The consumption of alcohol among recreational boaters is considered an important risk factor for fatalities and injuries among both operators and passengers. Using a database of all recreational vessels registered in Western Australia (WA), a sample of 500 adult boaters was recruited to participate in a telephone survey. The effects of demographic variables and boating characteristics upon the use of alcohol among recreational boaters on their last trip were explored using logistic regression. The odds of not having a drink were associated, after adjusting for age, with having completed a boating education course and with carrying children less than 12 years on board. The use of alcohol was not found to be prevalent among WA recreational boat owners. Based on these findings, it is recommended that efforts to decrease boating-related incidents, such as through education and legislation measures, be monitored over time to determine the effects of these strategies upon safety behaviours.

Differentiation of competitive vs. non-competitive mechanisms mediating disruption of moth sexual communication by point sources of sex pheromone (part 2): Case studies.

Eleven out of 13 disruption profiles (plots of dispenser density vs. male catch) from moth sex pheromone literature were consistent with a competitive-attraction mechanism, in which dispensers attract males and thereby divert them from females. Mean dispenser activity (D(a)) across all competitive-attraction cases was 0.04 +/- 0.06 (SD); values ranged from 0.0005 for a tiny laminated flake dispenser of racemic disparlure targeting gypsy moth to 0.2 for polyethylene tube dispensers used against lightbrown apple moth. A dispenser application activity (D(Aa)) can be calculated by multiplying D(a) by the number of such dispensers applied per hectare of crop. The highest dispenser application activity (D(Aa)) values approached 200 and corresponded to >99% inhibition of catches of male moths in monitoring traps. Relative to the D(Aa) scale, % inhibition of catches of male moths compressed and obscured large differences in D(Aa) when % disruption exceeded 90%. For cases of competitive attraction, these two efficacy scales can be interconverted by using the formula: D(Aa) approximately = 100/(100 minus % disruption). When disruptive point sources of pheromone were directly observed, male moths were seen approaching pheromone dispensers whose disruption profiles matched competitive attraction. Two cases fit non-competitive disruption mechanisms, which include camouflage, desensitization (adaptation and/or habituation), and sensory imbalance. In these cases, pheromone was released at rates higher than for cases of disruption by competitive attraction. Practical ramifications of the finding that competitive attraction appears to be the prevalent mechanism for moth mating disruption by pheromone point sources are listed. We believe that the congruence of diverse sets of mating disruption field data with explicit a priori predictions validates competitive-attraction theory. The analytical tools and principles governing competitive attraction that were uncovered during this study of mating disruption of moths should be generally applicable to competitive-attraction phenomena.

Differentiation of competitive vs. non-competitive mechanisms mediating disruption of moth sexual communication by point sources of sex pheromone (part I): Theory.

This study establishes a theoretical framework for differentiating among possible behavioral mechanisms whereby sexual communication of moths is disrupted in crops treated with point sources of pheromone. The major mechanisms recognized in the mating disruption literature fall into two main categories: competitive (competitive attraction = false-plume-following) and non-competitive (camouflage, desensitization, and sensory imbalance). Each disruption mechanism has been precisely defined verbally, and then the distinguishing characteristics of the two categories were defined mathematically. The sets of predictions associated with each category were visualized by graphical plots of mathematical simulations. Profiles of simulated male visitation rates to pheromone-baited traps deployed in pheromone-treated crops were graphed against density of pheromone dispensers by using various types of axes. Key traits of non-competitive attraction are as follows: concave profiles on untransformed axes, with an asymptotic approach to zero catch of male moths in traps; a straight line with positive slope when 1/catch is plotted against dispenser density (Miller-Gut plot); and a straight line with negative slope when catch is plotted against dispenser density * catch (Miller-de Lame plot). Key traits of non-competitive disruption profiles include: an initial linear disruption profile on untransformed axes; a concave Miller-Gut plot; and a recurving Miller-de Lame plot. These differences in profiles provide a basis for distinguishing competitive from non-competitive mechanisms when analyzing disruption profiles from field experiments. Slopes and intercepts of these secondary plots can also reveal both male and female moth densities, if the relative attractiveness of traps, females, and dispensers is known. The absolute value of the slope of the Miller-de Lame plot is a measure of each dispenser's activity (D(a)) for suppressing catch of male moths in traps. An application activity (D(Aa)) for a given dispenser can be calculated by multiplying D(a) by the number of such dispensers applied per hectare of crop.

Linked selected and neutral loci in heterogeneous environments.

We analyze a system of ordinary differential equations modeling haplotype frequencies at a physically linked pair of loci, one selected and one neutral, in a population consisting of two demes with divergent selection regimes. The system is singularly perturbed, with the migration rate m between the demes serving as a small parameter. We use geometric singular perturbation theory to show that when m is sufficiently small, each solution not initially fixed for the same selected allele in both demes approaches one of a 1-dimensional continuum of equilibria. We then obtain asymptotic expansions of the solutions and show their validity on arbitrarily long finite time intervals. From these expansions we obtain formulas for the transient dynamics of F(ST) (a measure of population structure) at both loci, as well as for the rate of genotyping error if the allelic state at the selected locus is inferred from that at the neutral (marker) locus. We examine two cases in detail, one modeling two populations in secondary contact after a period of evolution in allopatry, and the other modeling the origination and spread of a resistance allele.

Mechanized applicator for large-scale field deployment of paraffin-wax dispensers of pheromone for mating disruption in tree fruit.

A tractor-mounted mechanized applicator was developed for large-scale deployment of paraffin-wax dispensers of pheromone for mating disruption of oriental fruit moth, Grapholita molesta (Busck). The wax formulation was mostly water and emulsified paraffin wax containing 5% (by weight) pheromone [93:6:1 blend of (Z)-8-dodecen-1-yl-acetate:(E)-8-dodecen-1-yl-acetate: (Z)-8-dodecen-1-ol]. Ten milliliters of wax was applied per tree as approximately 160 deposits (0.04 ml of wax per drop). An average of 23 min was required to treat 1 ha of crop. Disruption efficacy of mechanically applied wax was measured relative to an untreated control in replicated 0.4-ha blocks within a recently abandoned apple orchard. From 6 May to 27 June, 100% disruption of tethered virgin females and 97% inhibition of pheromone traps was achieved for 52 d with two applications of wax. However, during mid- to late summer (July-August), this level of efficacy was maintained for only approximately 1 wk after each of two applications. Higher temperatures later in the season may have accounted for abbreviated efficacy of the applied small drops. Mechanically applied paraffin-wax technology may increase adoption of mating disruption given that a higher level of efficacy was achieved despite deploying less active ingredient per hectare relative to that used with reservoir dispensers. The savings in labor by not requiring hand application of reservoir dispensers could be directed toward cost of machinery. However, the short duration of efficacy obtained with the current wax formulation and mechanical applicator is judged uneconomical given the eight or more applications that would have been required for high-performance disruption over the full season. Larger drops with lower surface area-to-volume ratios are expected to prolong pheromone release for extended efficacy and desirable overall economics.

Higher densities of distributed pheromone sources provide disruption of codling moth (Lepidoptera: Tortricidae) superior to that of lower densities of clumped sources.

Field experiments quantified the effect of synthetic pheromone release-site density and distribution on 1) orientational disruption of male codling moths, Cydia pomonella (L.) (Lepidoptera: Tortricidae), to pheromone-baited traps; and 2) fruit injury. A clustering test varied pheromone release-site density from 0 to 1,000 Isomate-C Plus dispensers per ha while maintaining the total number of dispensers at 1,000. Percentage of orientational disruption of pheromone-baited traps increased significantly as a function of increasing density of release sites. Fruit injury decreased as the density of release sites increased and was lowest in plots treated with Isomate-C Plus dispensers distributed as 1,000 point sources per ha. We also manipulated point source density of 0.1-ml paraffin-wax drops containing 5% codlemone [(E,E)-8,10-dodecadien-1-ol], and thus the total amount of pheromone deployed per hectare. The percentage of disruption of traps baited with either 1.0- or 0.1-mg codlemone lures increased with increasing density of wax drops deployed. Both trapping and field observations confirmed that wax drops were attractive to male codling moths, suggesting that disruption was mediated by competitive attraction. Development of dispensers that can be mechanically applied at high densities has potential to improve the efficacy and economics of codling moth disruption at high population densities.

Orientational behaviors and EAG responses of male codling moth after exposure to synthetic sex pheromone from various dispensers.

The effect of brief pheromone exposures on responses of codling moth (Cydia pomonella L.) males was tested by flight-tunnel and electroantennogram (EAG) studies. Males were preexposed to pheromone for up to 3 min as they sat in release cages or for shorter times (a few seconds to several min) upon initiating flights or orienting in plumes. Brief exposures to Isomate-C Plus dispensers nearly eliminated moth orientations to 0.1 mg codlemone [(E,E)-8,10-dodecadien-1-ol] and 0.1 mg three-component (codlemone/12OH/14OH, 100:20:5 ratio) lures 15 min later. However, there was no associated change in EAG responses between preexposed and control moths. Behavioral responses of Isomate-C Plus-exposed males were normal 24 hr following exposure. The reduced sexual responsiveness observed following exposure to Isomate dispensers appeared to be associated with an elevation of response threshold. Brief preexposure to 0.1 mg codlemone and three-component lures also reduced orientational behavior of males 15 min later, but to a lesser degree than when preexposed to Isomate-C Plus dispensers. Male behavior following preexposure to a 0.1 mg codlemone/pear ester [(2E,4Z)-2,4-decadienoate] lure (1:1 ratio) was no different from exposure to codlemone only. Orientational disruption in plots treated with 10 dispensers of Isomate-C Plus per tree was 88.3 and 95.9% for 1.0 and 0.1 mg codlemone lures, respectively. Some males did orient to 0.1 mg codlemone lures so we caution that flight-tunnel experiments on preexposure may overestimate the actual pheromone exposure dosage received by feral moths in treated orchards. Importantly, this work documents that a portion of feral males within a population has the capacity to overcome communicational disruption by high densities of Isomate-C Plus dispensers.

A finite locus effect diffusion model for the evolution of a quantitative trait.

A diffusion model is constructed for the joint distribution of absolute locus effect sizes and allele frequencies for loci contributing to an additive quantitative trait under selection in a haploid, panmictic population. The model is designed to approximate a discrete model exactly in the limit as both population size and the number of loci affecting the trait tend to infinity. For the case when all loci have the same absolute effect size, formal multiple-timescale asymptotics are used to predict the long-time response of the population trait mean to selection. For the case where loci can take on either of two distinct effect sizes, not necessarily with equal probability, numerical solutions of the system indicate that response to selection of a quantitative trait is insensitive to the variability of the distribution of effect sizes when mutation is negligible.