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Reduction in size and fecundity of the autumnal moth, Epirrita autumnata, in the increase phase of a population cycle.

Increasing fecundity with increasing density has been observed for many cyclic herbivore populations, including some forest Lepidoptera. We monitored population density, body size and reproductive capacity of the cyclic lepidopteran, the autumnal moth (Epirrita autumnata, Geometridae), from the early increase phase to the devastating outbreak density in northernmost Norway. Larval density of the species increased exponentially from 1998 to 2002 and remained at the outbreak level also in 2003. Within the same period, the body size and fecundity of individuals reduced as analysed from several parallel datasets on larvae, pupae and adults. In another study area in northernmost Finland, the density increase of the autumnal moth was moderate only, and true outbreak density was not attained during the study. Despite that, a reduction was again detected in the size and fecundity of individuals. Possible factors responsible for the reduced size and fecundity of individuals in the Norwegian population were quantitative shortage of foliage, rapid and delayed inducible resistances of the host, mountain birch (Betula pubescens ssp. czerepanovii), as well as crowding-induced responses of larvae. These factors likely acted in concert, although non-delayed responses to the density were emphasized. Our findings did not support the hypotheses of climatic release, inducible susceptibility of the host tree and mast depression (i.e. lowered chemical defence of the host tree after its mast seeding) as promoters of the fecundity-based density increase of the autumnal moth, since the reduced fecundity in relation to increased density was strongly against the predictions of these hypotheses. Therefore, we suggest that the density increase of autumnal moth populations is promoted by high survival rather than exceptionally high fecundity.

Animals↗

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.

Animals↗

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.

Animal Communication↗

Detection and characterisation of three novel species of reovirus (Reoviridae), isolated from geographically separate populations of the winter moth Operophtera brumata (Lepidoptera: Geometridae) on Orkney.

Geographically separate populations of winter moth (Operophtera brumata L.) were sampled in heather habitats on the Orkney Isles in order to investigate the prevalence of virus pathogens. Reoviruses were isolated in 11 of the 13 winter moth populations sampled, with 3 novel species being detected. Two species of Cypoviridae (CPV) were isolated, Operophtera brumata CPV18 and O. brumata CPV19, with one host population suffering 46% infection prevalence of OpbuCPV19. A third virus, O. brumata Reovirus (OpbuRV), was isolated from both winter moth and a hymenopteran parasitoid wasp, Phobocampe tempestiva, which is abundant in these populations. This was identified as a non-occluded reovirus, which was clearly able to infect and persist in both the lepidopteran and the hymenopteran host. The genomes of the three viruses were characterised using gel electrophoresis and the virus structure was investigated using transmission electron microscopy. The relationship of these viruses with a baculovirus that also infects winter moth, OpbuNPV, was investigated, as well as the association of OpbuRV with P. tempestiva. The detection of such viruses is discussed with reference to studies of similar viruses in other lepidopteran and hymenopteran host systems.

Animals↗

Chemical constraints on the evolution of olfactory communication channels of moths.

It is estimated there are over 100,000 moth species (Lepidoptera) that produce sex pheromones comprising communication channels used in specific-mate-recognition systems (SMRS) involved in pre-zygotic mating isolation and speciation. About 1572 moth species have been found to use 377 pheromone components, the majority being alcohols, aldehydes or acetate esters of olefinic chains of 10-22 carbons. Since there are limited numbers of possible unsaturated (double bonded) E- and Z-configured isomers of these chains, there may be constraints on incipient species evolving new pheromone components, especially if they are utilized by existing species. Here I develop algorithms that count and name all possible structures of chains with or without a functional group. The results show that for acetate esters there are only seven or nine monounsaturated isomers of six or seven carbon chains, respectively, suggesting use of these compounds could limit the number of communication channels available for radiation of new species (no moths use these short chains). For commonly used 14-carbon chains with an acetate ester functional group, and 1-3 unsaturations, there are 1039 isomers. A total of 2,096,883 isomers were counted from all multiply unsaturated 10-22-carbon chain acetate esters. The number of possible signals quickly extends into millions when considering pheromone blends of 2-4 components used by most species. There should be little chemical constraint on evolution of new communication channels based on compounds of 10 or more carbons, even for closely related species (e.g. ermine moths, Yponomeuta).

Acetates↗

Appetitive flight patterns of male Agrotis segetum moths over landscape scales.

An analysis is presented of the first harmonic radar studies of pheromone-plume locating flights of male Agrotis segetum moths over distances of up to 500 m. Upon release most moths flew in a direction having a downwind component. The first significant changes in flight orientations occur in the immediate vicinity of a pheromone source. Moths that were initially flying downwind change course and start flying crosswind whilst those that initially flew crosswind change course and start flying upwind. It is shown that such behaviour is consistent with the adoption of an effective plume-location strategy, and conditions are identified when downwind flights would be more advantageous than crosswind ones. Additionally, some of the complex flight patterns that can arise at later times are shown to be compatible with the adoption of an optimal biased scale-free (Lévy-flight) searching strategy. It is found that disruptive doses of sex pheromone can have a marked influence upon male moth flight patterns.

Animals↗

Complete nucleotide sequence of the vitellogenin mRNA from the gypsy moth: novel arrangement of the subunit encoding regions.

Primary structure analysis and location of introns suggests evolutionary relatedness among vitellogenin (Vg) genes from vertebrates and invertebrates, including insects. We have determined the complete nucleotide sequence of the gypsy moth VgmRNA, which shows that its structure is significantly different from VgmRNAs in other systems. The nucleotide sequence was determined using overlapping cDNA fragments generated from RACE reactions and rTh polymerase-mediated PCR. The VgmRNA is 5579 nucleotides long and codes for both the large and small subunits. However, the arrangement of the subunit encoding regions in the gypsy moth VgmRNA is opposite of what has been observed in other systems. Gypsy moth Vg gene is the first reported example of a Vg gene where the 5'-terminal region codes for the large subunit and the 3'-terminal region for the small subunit. Also, the sequence near the junction of subunits was significantly different from those found in other insects. This may be responsible for the relatively more stable precursor of Vg subunits found in the gypsy moth hemolymph. It is not clear where this divergence in the structure of Vg gene occurred during evolution, since the Vg gene of another lepidopteran, Bombyx mori, conforms to the structure of those in vertebrates and other invertebrates.

Amino Acid Sequence↗

Discrimination of pheromone enantiomers by two pheromone binding proteins from the gypsy moth Lymantria dispar.

The gypsy moth, Lymantria dispar, uses (7R, 8S)-cis-2-methyl-7, 8-epoxyoctadecane, (+)-disparlure, as a sex pheromone. The (-) enantiomer of the pheromone is a strong behavioral antagonist. Specialized sensory hairs, sensillae, on the antennae of male moths detect the pheromone. Once the pheromone enters a sensillum, the very abundant pheromone binding protein (PBP) transports the odorant to the sensory neuron. We have expressed the two PBPs found in gypsy moth antennae, PBP1 and PBP2, and we have studied the affinity of these recombinant PBPs for the enantiomers of disparlure. To study pheromone binding under equilibrium conditions, we developed and validated a binding assay. We have addressed the two major problems with hydrophobic ligands in aqueous solution: (1) concentration-dependent adsorption of the ligand on vial surfaces and (2) separation of the protein-bound ligand from the material remaining free in solution. We used this assay to demonstrate for the first time that pheromone binding to PBP is reversible and that the two PBPs from L. dispar differ in their enantiomer binding preference. PBP1 has a higher affinity for the (-) enantiomer, while PBP2 has a higher affinity for the (+) enantiomer. The PBP from the wild silk moth, Antheraea polyphemus (Apol-3) bound the disparlure enantiomers more weakly than either of the L. dispar PBPs, but Apol-3 was also able to discriminate the enantiomers. We have observed extensive aggregation of both L. dispar PBPs and an increase in pheromone binding at high (>2 microM) PBP concentrations. We present a model of disparlure binding to the two PBPs.

Alkanes↗

Syntheses of (Z,E)-5,7-dodecadienol and (E,Z)-10,12-hexadecadienol, lepidoptera pheromone components, via zinc reduction of enyne precursors. test of pheromone efficacy against the Siberian moth.

Efficient syntheses of (Z,E)-5,7-dodecadienol, a pheromone component of the Siberian moth, Dendrolimus superans sibiricus, and (E,Z)-10,12-hexadecadienol, a pheromone component of various Lepidoptera pheromones, were accomplished by cis reduction of the corresponding enynols with activated zinc. The most energetic reagent was zinc galvanized with copper and silver (Zn/Cu/Ag) that achieved rapid and high-yield reduction in methanol-water. The stereoselectivity of semi-hydrogenation was > or =98%. A process whereby zinc dust was continuously activated throughout the reduction with an acid was also satisfactory (95-98% cis). Field evaluation of the 1:1 mixture of (Z,E)-5,7-dodecadienol and (Z,E)-5,7-dodecadienal with the Siberian moth in Russia showed that the rubber septa pretreated with compound and stored at -80 degrees C were as effective as freshly treated septa. Moth responses to septa aged in open air indicated that lure effectiveness declined significantly after 2 weeks of aging. Thus, if rubber septa are used as pheromone dispensers in Siberian moth traps monitoring, they should be replaced biweekly with fresh septa for optimal trap effectiveness.

Animals↗

Volatiles associated with preferred and nonpreferred hosts of the Nantucket pine tip moth, Rhyacionia frustrana.

Ovipositing female Nantucket pine tip moth, Rhyacionia frustrana, prefer loblolly pine, Pinus taeda L., to slash pine, Pinus elliottii Engelm, except during the first spring following planting of seedlings. Host discrimination by R. frustrana increases as seedlings develop, suggesting that changes in the chemical composition of seedlings may mediate the moth's host preferences. Volatile compounds from slash and loblolly pine seedlings were collected using solid-phase microextraction (SPME) during the first year following planting. Four collection periods coincided with adult emergence and oviposition during each of four annual generations of R. frustrana in the Georgia Coastal Plain. Infestation of slash pine peaked during the second tip moth generation and was similar to the loblolly pine infestation level. By the fourth tip moth generation, slash pine infestation levels had declined and diverged considerably from those of loblolly pine. Significant differences in relative quantities of beta-pinene, alpha-phellandrene, limonene, beta-phellandrene, bornyl acetate, beta-caryophyllene, and an unidentified sesquiterpene occurred between slash and loblolly pine during the fourth generation. However, no strong correlation was observed between any individual compound and host damage that could readily explain the temporal changes in R. frustrana host preference. Gas chromatographic-electroantennographic detection (GC-EAD) analyses of standards identified 19 different seedling-associated compounds that elicited antennal responses from R. frustrana females, indicating that a blend of terpenoids may mediate host discrimination.

Animals↗

Adaptation of antennal neurons in moths is associated with cessation of pheromone-mediated upwind flight.

A wind-borne plume of sex pheromone from a female moth or a synthetic source has a fine, filamentous structure that creates steep and rapid fluctuations in concentration for a male moth flying up the plume's axis. The firing rates from single antennal neurons on Agrotis segetum antennae decreased to nearly zero within seconds after the antennae were placed in a pheromone plume 70 cm downwind of a high-concentration source known from previous studies to cause in-flight arrestment of upwind progress. In a separate experiment, the fluctuating output from chilled neurons on Grapholita molesta antennae became attenuated in response to repetitive, experimentally delivered pheromone pulses. The attenuation was correlated with a previously reported higher percentage of in-flight arrestment exhibited by moths flying at cooler compared to warmer temperatures. These results indicate that two peripheral processes related to excessive concentration, complete adaptation of antennal neurons, or merely the attenuation of fluctuations in burst frequency, are important determinants of when upwind progress by a moth flying in a pheromone plume stops and changes to station keeping. Also, adaptation and attenuation may affect the sensation of blend quality by preferentially affecting cells sensitive to the most abundant components in airborne pheromone blends.

Action Potentials↗

Dose-morality and large-scale studies for controlling codling moth (Lepidoptera: Tortricidae) eggs on 'd'Agen' plums by using methyl bromide.

Codling moth, Cydia pomonella (L.), eggs on 'd'Agen' cultivar of plums, Prunus salicina Lindl., were treated with methyl bromide to determine if a quarantine treatment could be developed so that the plums could be exported to Japan from the United States. Small-scale tests consisted of treating codling moth eggs on plums with various doses of methyl bromide at 20 degrees C for 2 h. Small-scale tests showed that 0- to 24-h-old eggs of codling moth on the plums were controlled by doses > 22.5 g/m3. Because testing showed that 48 g/m3 had no adverse effect on the quality of plums, this dose was chosen for large-scale testing to establish the quarantine dose. Large-scale tests consisted of treating plums at 18.5 degrees C for 2 h using methyl bromide at a dose of 48 mg/liter. Large-scale tests showed that the dose of 48 g/m3 killed all 0- to 24-h-old codling moth eggs exposed on plums in packing cartons without affecting the quality of the plums.

Animals↗

Resistance in sunflower and interaction with Bacillus thuringiensis for control of banded sunflower moth (Lepidoptera: Tortricidae).

Four sunflower accessions were compared with a susceptible check, hybrid '894', in the greenhouse to determine their resistance to the banded sunflower moth, Cochylis hospes Walsingham, and their interaction with Bacillus thuringiensis Berliner variety kurstaki. Antibiosis, expressed as lower larval weight, was detected in all of the accessions. In addition to being antibiotic, sunflower accession Ames 3291 was antixenotic to banded sunflower moth oviposition and exhibited an additional impact on larval weight when B. thuringiensis was applied. By itself, B. thuringiensis provided better control of banded sunflower moth than the resistance tested. However, banded sunflower moth-resistant sunflower would be a good option when B. thuringiensis or another insecticide is not applied, and it may prevent the economic threshold from being reached.

Animals↗

Viability of Claviceps africana spores ingested by adult corn earworm moths, Helicoverpa zea (Boddie) (Lepidoptera: Noctuidae).

A study was conducted in College Station, TX, to determine the viability of Claviceps africana spores in the digestive tract of adult corn earworm moths, Helicoverpa zea (Boddie). Both sexes were exposed to ergot-infected sorghum panicles for 30 min, and spores were recovered from excreta of the moths at 24-, 48-, and 72-h intervals after feeding. Recovered spores were quantified, and viability was determined by the germination rate of macroconidia. Nearly a 100-fold greater concentration of spores was recovered from female excreta at the three time intervals compared with male excreta. Concentration of spores in female and male excreta was greatest at 24 h, with a significant reduction at the later time intervals. Spore germination rates for both sexes were greater at 24 h, with survival being significantly reduced at the 72-h interval. Spores in female excreta survived longer than those from male excreta. Spore survival over time was significantly reduced in male excreta. Spore concentration and survival were greater from female excreta, which is key, because egg-laying activities on sorghum panicles intensify during flowering, and this source of ergot spores could contribute to the spread of the disease. This study demonstrates that corn earworm moths can internally carry viable ergot spores for several days and can act as primary dispersal agents for the fungus. This is important because contaminated moths migrating from areas in Mexico and southern Texas where ergot is endemic could transmit and spread the disease to other sorghum growing regions of the United States.

Animals↗

Inheritance of a temperature-modified phenotype of the short antennae (sa) mutation in a moth, Ephestia kuehniella (Lepidoptera: Pyralidae).

The autosomal recessive mutation short antennae (sa) causes considerable shortening of antennae in male and female Mediterranean flour moths (Ephestia kuehniella Zeller). However, the sa phenotype can be suppressed by several physical factors, making sa moths indistinguishable from wild-type moths (sa(WT)). This can be done by subjecting larva and pupa to a higher temperature (25 degrees C), to lithium ions, or to an alternate electric field. The first half of pupal development was found to be the sensitive period for the sa(WT) phenotype. The sa(WT) phenotype is stable and cannot be reverted to the original sa type by physical or chemical factors. The sa(WT) phenotype is transmitted to future generations. When crossed with typical sa moths, the sa(WT) phenotype is inherited either as a dominant character if carried by males or a semidominant character if carried by females. We compared proteins of the ejaculate, accessory gland secretions, and spermatophore in sa, sa(WT), and wild-type males and found considerable differences between sperm proteins of sa(WT), sa, and wild-type males. The sa(WT) phenotype influences the mating success of males: sa(WT) males mated successfully with any females, whereas typical sa males were less successful in mating and then mainly with females of the same phenotype.

Animals↗

Patterns of genetic structure among populations of an oligophagous pollinating yucca moth (Tegeticula yuccasella).

Plant-insect associations have served as models for investigations of coevolution and the influence of biotic interactions on diversification. The pollination association between yuccas and yucca moths is a classic example of an obligate mutualism often suggested to have been affected by coevolution. Recent work has shown high host specificity in pollinating yucca moths, and here we use Tegeticula yuccasella, the species with the widest diet breadth, to ask how host specificity and isolation by distance contribute to specialization. Isolation by distance at a regional scale was observed in nucleotide variation within the mitochondrial gene cytochrome oxidase I (COI) (r =.294; P =.003). Host-related genetic structure (F(ct) = 0.08) was found to be slightly lower than the level of structure observed between eastern and western moth populations (F(ct) = 0.096). However, 56% of the COI haplotypes sampled from moths on Yucca filamentosa mapped to a host-specific clade in the haplotype network. Taken together, these results suggest that differentiation among T. yuccasella populations on alternative hosts is slight, but gene flow is influenced by both host association and geographic distance.

Animals↗

Persistence of bat defence reactions in high Arctic moths (Lepidoptera).

We investigated the bat defence reactions of three species of moths (Gynaephora groenlandica, Gynaephora rossi (Lymantriidae) and Psychophora sabini (Geometridae)) in the Canadian Arctic archipelago. Since these moths inhabit the Arctic tundra and, therefore, are most probably spatially isolated from bats, their hearing and associated defensive reactions are probably useless and would therefore be expected to disappear with ongoing adaptation to Arctic conditions. When exposed to bat-like ultrasound (26 kHz and 110 dB sound pressure level root mean square at 1 m) flying male Gynaephora spp. always reacted defensively by rapidly reversing their flight course. They could hear the sound and reacted at least 15-25 m away. Psychophora sabini walking on a surface froze at distances of at least 5-7 m from the sound source. However, two out of three individuals of this species (all males) did not respond in any way to the sound while in flight. Hence, we found evidence of degeneration of bat defence reactions, i.e. adaptation to the bat-free environment, in P. sabini but not in Gynaephora spp. Some Arctic moths (Gynaephora spp.) still possess defensive reactions against bats, possibly because the selection pressure for the loss of the trait is such that it declines only very slowly (perhaps by genetic drift; and there may not have been enough time for the trait to disappear. One possible reason may be that Arctic moths have long generation times.

Animals↗

Insect odour perception: recognition of odour components by flower foraging moths.

Odours emitted by flowers are complex blends of volatile compounds. These odours are learnt by flower-visiting insect species, improving their recognition of rewarding flowers and thus foraging efficiency. We investigated the flexibility of floral odour learning by testing whether adult moths recognize single compounds common to flowers on which they forage. Dual choice preference tests on Helicoverpa armigera moths allowed free flying moths to forage on one of three flower species; Argyranthemum frutescens (federation daisy), Cajanus cajan (pigeonpea) or Nicotiana tabacum (tobacco). Results showed that, (i) a benzenoid (phenylacetaldehyde) and a monoterpene (linalool) were subsequently recognized after visits to flowers that emitted these volatile constituents, (ii) in a preference test, other monoterpenes in the flowers' odour did not affect the moths' ability to recognize the monoterpene linalool and (iii) relative preferences for two volatiles changed after foraging experience on a single flower species that emitted both volatiles. The importance of using free flying insects and real flowers to understand the mechanisms involved in floral odour learning in nature are discussed in the context of our findings.

Acetaldehyde↗