Pheromone-mediated upwind flight of male gypsy moths, Lymantria dispar, in a forest

Abstract Lymantria dispar L. males flying upwind in a pheromone plume in a forest were video-recorded at 2.5, 10 and 20 m from the source of pheromone. Males flew slower and steered more across the wind as they approached the source. In concert, their ground speed decreased and track angles increased. In contrast to these changes, their drift angles were fairly constant and the transverse component of image flow, above and/or below the moths eyes, showed almost no change. The inter-turn duration (time between sequential turns), a temporal aspect of the male flight manoeuvres, showed a consistent but relatively small increase as the distance from the source increased. The flight tracks narrowed as the males approached close (2.5 m) to the source. Because of unpredicted correlations between physical variables (i.e. temperature, wind velocity) and the distance from the source, we used principal components analysis to generate a set of completely independent variables. Greater than 90% of the variability in the data could be explained by four principal factors which corresponded well with known relationships in the flight manoeuvres. All four of these factors showed a significant regression against distance to the source. Although uncontrolled factors such as temperature and wind velocity may have contributed to changes in flight behaviour, recent data indicate that, in addition to concentration, certain temporal and spatial characteristics (i.e. burst period, burst return period) of plumes in wind vary systematically with distance from the source. We propose that L.dispar males might adjust their flight manoeuvres in response to these changes.     

Host tree influences on the dispersal of first instar gypsy moths, Lymantria dispar (L.)

Abstract. 1. In laboratory tests, first instar gypsy moths attempted dispersal more frequency when exposed to less acceptable foliage.
2. First instars from small eggs attempted dispersal less frequently than larvae from large eggs when exposed to foliage from highly acceptable or marginally acceptable hosts. Dispersal rates of larvae from medium sized eggs were intermediate.
3. These results (1–2) confirm and expand upon the findings of Capinera & Barbosa (1976).
4. In the field, data on the relative densities of larvae on different host species support the conclusion that the frequency of dispersal attempts is inversely related to host acceptability.
5. The implications of these findings for the population dynamics of the gypsy moth are discussed.     

Pheromonotropic activity in the gypsy moth Lymantria dispar: evidence for a neuropeptide

The brain-suboesophageal ganglion complex of the gypsy moth, Lymantria dispar, contains pheromonotropic activity detectable using a Helicoverpa zea in vivo bioassay for pheromone-biosynthesis-activating neuropeptide. Pheromonotropic activity was detected as early as the third larval instar and was present throughout development and through day 6 post-eclosion. Activity in the adult is presumably associated with pheromone production, while it is speculated that larval activity may be related to melanization. Adult pheromonotropic activity is associated with a peptide of approximately 3.500 kDa. It is heat labile and only partially stable when incubated at 35°C or exposed to freeze-thawing. Isolation of L. dispar pheromonotropic factor should facilitate the elucidation of the mechanism of pheromone production in this insect pest.Abbreviations ED₅₀ dose at which one-half maximal response is observal - eq equivalent - MRCH melanization and reddish colorization hormone - MW molecular weight - PBAN pheromone biosynthesis activating neuropeptide - SOG suboesophageal ganglion - TFA trifluoroacetic acid - Z11-16: Ald (Z)-11-hexadecenal     

Visual position stabilization in the hummingbird hawk moth, Macroglossum stellatarum L. I. Behavioural analysis

Optomotor responses of freely flying hawk moths, Macroglossum stellatarum, were characterized while the animals were hovering in front of and feeding on a dummy flower. Compensatory translational and rotational movements of the hawk moth were elicited by vertical grating patterns moving horizontally, mimicking imposed rotational and translational displacements of the animal in the horizontal plane. Oscillatory translational and rotational pattern motion leads to compensatory responses that peak in the frequency range between 2 Hz and 4 Hz. The control systems mediating the translational and rotational components of the optomotor response do not seem to influence each other. The system mediating translational responses is more sensitive in the fronto-lateral part of the visual field than in the lateral part; the opposite is true for the rotational system. The sensitivity of the translational system does not change along the vertical, whereas the rotational system is much more sensitive to motion in the dorsal than in the ventral part of the visual field. These sensitivity gradients may reflect an adaptation to the specific requirements of position stabilization in front of flowers during feeding. Accepted: 13 August 1997     

Orientation of male gypsy moths,Lymantria Dispar (L.), to pheromone sources: The role of olfactory and visual cues

The role of olfaction and vision in the close-ranging flying and walking orientation of male gypsy moths, Lymantria dispar(L.), to females was studied in the forest and in the laboratory. In the forest, feral males found an isolated pheromone source as readily as one supplemented with female visual cues; dead, acetonerinsed females deployed without pheromone received virtually no visitations. In flight tunnel choice experiments using cylinders as surrogate trees and pheromone in different spatial configurations, visual attributes of the female did not influence either the males' choice of landing site or the efficiency with which they located the female. Rather, the presence of pheromone on the cylinder was necessary to elicit orientation as well as landing and walking on the cylinder. When a female visual model was placed in various positions around a pheromone source, walking males oriented primarily to the chemical stimulus. There were, however, indications that males would alter their walking paths in response to female visual cues over short distances (<5 cm), but only if they continued to receive pheromone stimulation. When visual and chemical cues were abruptly uncoupled by altering the trajectory of the pheromone plume, most males responded to the loss of the odor cue rather than to visual cues from the female. Temporal pheromone stimulation patterns affected male walking orientation. When stimulated by pheromone, males oriented toward the source; loss of the odor cue prompted an arearestricted local search characterized by primarily vertical and oblique movements with frequent reversals in direction. Presumably these maneuvers enhance the likelihood of recontacting the plume or serendipitously encountering the female. The apparent lack of visual response to the female is discussed in light of morphological and behavioral evidence suggesting that gypsy moths were formerly nocturnal.     

Visual position stabilization in the hummingbird hawk moth, Macroglossum stellatarum L. II. Electrophysiological analysis of neurons sensitive to wide-field image motion

Response properties of neurons in the cervical connectives of the hummingbird hawk moth, Macroglossum stellatarum L., were determined. All neurons described in this account respond directionally selectively to motion in large parts of the visual field of either eye. They respond maximally to bilateral stimulation, preferring either motion as induced on the eyes during translatory movements of the animal or when it turns around one of its body axes. Cells most sensitive to rotational motion either respond best to rotation of the patterns around the vertical axis of the animal or around its longitudinal body axis. Neurons most sensitive to translational pattern motion respond best to either simulated translations of the animal along its vertical or along an oblique axis. Most types of neurons respond tonically and do not habituate. The sensitivity to motion stimuli is not evenly distributed within the receptive field of any investigated neuron. Part of these neurons might play a role in visual position and course stabilization. Accepted: 13 August 1997     

Effects of pheromone plume structure and visual stimuli on the pheromone-modulated upwind flight of male gypsy moths (Lymantria dispar) in a Forest (Lepidoptera: Lymantriidae)

The pheromone-modulated upwind flight ofLymantria dispar males responding to different pheromone plume structures and visual stimuli designed to mimic trees was video recorded in a forest. Males flying upwind along pheromone plumes of similar structure generated tracks that were similar in appearance and quantitatively similar in almost all parameters measured, regardless of the experimentally manipulated visual stimuli associated with the pheromone source. Net velocities, ground speeds, and airspeeds of males flying in point-source plumes were slower than those of males flying in the wider, more diffuse plumes issuing from a cylindrical baffle. The mean track angle of males flying in plumes issuing from a point source was greater (oriented more across the wind) than that of males flying in plumes issuing from a transparent cylindrical baffle. Males flying in point-source plumes also turned more frequently and had narrower tracks overall than males responding to plumes from a cylindrical baffle. These data suggest thatL. dispar males orienting to pheromone sources (i.e., calling females) associated with visible vertical cylinders (i.e., trees) use predominantly olfactory cues to locate the source and that the structure of the pheromone plume markedly affects the flight orientation and the resultant track.     

Structure and function of antennal lobe neurons in the male turnip moth,Agrotis segetum (Lepidoptera: Noctuidae)

Interneurons with dendritic branches in the antennal lobe of the male turnip moth, Agrotis segetum (Schiff., Lepidoptera: Noctuidae), were investigated with intracellular recording and staining methods. Seventeen projection neurons that transmit information from the antennal lobe to higher centers in the brain displayed dendritic arbors in the male specific macroglomerular complex (MGC) and responded to chemical components of the female sex pheromone used in species-specific sexual communication. Most of the projection neurons responded to several of the pheromone components tested, and a precise correlation between the location of the dendritic arborization and the physiological response could not be demonstrated. One MGC-projection neuron fit the definition of blend specialist. It did not respond to the individual components of the behaviorally active pheromone blend, but showed a strong response to the components when combined in the species-specific blend. Some of the projection neurons also showed clear responses to phenylacetaldehyde, a flower-produced compound and/or to (E)-2-hexenal, a common green-leaf volatile. In eight neurons, the axonal projection could be followed to the calyces of the mushroom body, and subsequently to the inferior lateral protocerebrum.Four local interneurons were characterized both morphologically and physiologically. Each neuron arborized extensively throughout the antennal lobe, and each responded to one or several of the pheromone compounds, and/or to one or both of the plant-produced compounds. One of the local interneurons responded exclusively to the pheromone blend, but not to the individual components.Abbreviations AL antennal lobe - AN antennal nerve - CB cell body - E2H (E)-2-hexenal - IACT inner antennocerebral tract - ILPR inferior lateral protocerebrum - LH lateral horn of the protocerebrum - LN local interneuron - MB mushroom body - MGC macroglomerular complex - OACT outer antennocerebral tract - PAA phenylacetaldehyde - PN projection interneuron - RN receptor neuron - Z5-10:OAc (Z)-5-decenyl acetate - Z5-10:OH (Z)-5-decenol - Z5-12:OAc (Z)-5-dodecenyl acetate - Z7-12:OAc (Z)-7-dodecenyl acetate - Z9-14:OAc (Z)-9-tetradecenyl acetate     

Vertical transmission and overwintering of microsporidia in the gypsy moth, Lymantria dispar

Vertical transmission and the overwintering success of three different microsporidia infecting Lymantria dispar (Lepidoptera: Lymantriidae) larvae were investigated. Endoreticulatus schubergi, a midgut pathogen, was transmitted to offspring via female and male via the egg chorion (transovum transmission). Between 8% and 29% of the emerging larvae became infected. No spores of E. schubergi were found in surface-washed eggs. Nosema lymantriae, a microsporidium that causes systemic infections, was transovarially transmitted. Between 35% and 72% of the progeny were infected. Vairimorpha disparis, a fat body pathogen, was not vertically transmitted. The infectivity of spores that overwintered in cadavers of infected L. dispar varied by species, placement in the environment, and weather conditions. Spores of E. schubergi were still infective after an eight month exposure period of cadavers on the ground. Spores of N. lymantriae and V. disparis remained highly infective only when cadavers overwintered under a more or less continuous snow cover for four months.     

Iridoid glycosides and insect feeding preferences: gypsy moths (Lymantria dispar, Lymantriidae) and buckeyes (Junonia coenia, Nymphalidae)

Abstract. 1. The effect of increasing concentrations of an iridoid glycoside, catalposide, was tested on a generalist and an adapted specialist lepidopteran, using artificial diets.
2. Two strains of the gypsy moth, Lymantria dispar (L.) (Lymantriidae), were tested. Larvae of a wild strain showed a negative dose-dependent response to increasing concentrations of catalposide (up to 7.2% dry weight), while a laboratory strain showed no such pattern. Choice tests using the laboratory strain showed that these larvae often preferred diets relatively low in catalposide over those with higher concentrations.
3. In contrast, larvae of the iridoid glycoside specialist, the buckeye, Junonia coenia Hübner (Nymphalidae), showed a positive dose-dependent response and grew better on diets with higher doses of catalposide.
4. We tested whether the improved growth of J. coenia at higher concentrations of catalposide may have been due to extra glucose yielded by hydrolysis of catalposide. Larvae were fed increasing doses of glucose equivalent to the amount potentially obtained by hydrolysing the quantities of catalposide used in the previous experiment. However, added glucose did not effect growth, suggesting that increased growth at higher catalposide concentrations was due to increased feeding rates.     

Phenology of Lymantria dispar (Lepidoptera: Lymantriidae), male flight and the effect of moth dispersal in heterogeneous landscapes

 A model of Lymantria dispar development was assembled from the published literature and used to predict the period of male moth flight in the United States. Model predictions were compared with observations made with pheromone traps in several locations throughout the United States but especially in Virginia, West Virginia and North Carolina between 1995 and 1996. The model was found to provide accurate and unbiased forecasts of the dates of 5%, 50% and 95% cumulative trap catch, particularly at lower elevations. In areas of high topographic diversity (such as West Virginia), deviations between model output and observations were minimized by basing predictions of 5% and 50% cumulative catch on minimum elevation within neighborhoods of 25–81 km². This model of L. dispar male flight phenology can be used to time the deployment and retrieval of pheromone traps in intensive or extensive monitoring programs. However, a better understanding of moth movement is needed to fully explain the patterns of local trap catch. Received: 9 October 1997/Accepted: 8 December 1997     

Characterisation of three Alphabaculovirus isolates from the gypsy moth,Lymantria dispar dispar (Lepidoptera: Erebidae), in Turkey

In this study, Lymantria dispar dispar larvae, collected from three different localities in Turkey, were examined for the presence of inclusion bodies under phase contrast and electron microscopes. Inclusion bodies from infected larvae were subjected to polymerase chain reaction using the conserved primers for polyhedrin (polh), late expression factor 8 (lef-8) and late expression factor 9 (lef-9) genes. Sequence analysis confirmed that larvae collected from the three different localities contained multiple nucleopolyhedrosis viruses (MNPVs). These isolates were designated LdMNPV-T1, LdMNPV-T2 and LdMNPV-T3. Phylogenetic analyses of these isolates were performed using target genes polh, lef-8 and lef-9. Restriction endonuclease analysis of the three geographic isolates with EcoRI and PstI enzymes demonstrated some differences existed among the isolates. According to the EcoRI profile, the mean estimated size for the complete genome of each isolate (LdMNPV-T1, LdMNPV-T2 and LdMNPV-T3) was calculated to be approximately 170, 153 and 170?kb, respectively. Insecticidal activities of each isolate were tested on L. d. dispar larvae using four different viral concentrations between 10³ and 10⁶?OBs/ml. Results showed that the mortalities for LdMNPV-T1, -T2 and -T3 ranged between 13–53%, 47–100% and 46–93%, respectively. The LC₅₀ and LC₉₅ values of LdMNPV-T2 were not significantly different from the respective corresponding values of the other two isolates. However, isolate LdMNPV-T2 killed larvae with a LC₅₀ value that was lower than the other two isolates. Our results suggested there are promising LdMNPV isolates in Turkey that can be used for microbial control of L. d. dispar larvae.     

Long-term laboratory evaluation ofRogas lymantriae: A braconid endoparasite of the gypsy moth,Lymantria dispar

Rogas lymantriae Watanabe was reared onLymantria dispar (L.) for 125 generations in the laboratory following importation from Japan. No deleterious effects of colonization were observed and one measured parameter, successful parasite emergence, significantly improved over time. Decreased density of the host or conspecific female parasites resulted in higher levels of parasitism and numbers of females in the next generation. During 3 different years, colony production was significantly increased with little difficulty to provide parasites for inoculative releases in the field.      

Ecdysone 20-hydroxylation and 3-epimerization in larvae of the gypsy moth,Lymantria dispar L.: tissue distribution and developmental changes

The potential for ecdysone metabolism was determined for various larval tissues of the gypsy moth, Lymantria dispar. Homogenates of fat body, midguts, and Malpighian tubules, taken on different days during the second half of the fifth instar, were incubated with [(3)H]ecdysone, and the products were analyzed by reversed-phase and normal-phase HPLC. All tissues showed conversion to 20-hydroxyecdysone, and midguts also produced 3-epiecdysone. Ecdysone 20-monooxygenase (E20MO) activity in the fat body increased from a low level on day 5 to a peak on day 11, coinciding with the peak in the hemolymph ecdysteroid titer on the penultimate day of the instar. Midguts and Malpighian tubules showed E20MO activity only during the last 3 or 4days of the instar, with the highest activity also occurring on the penultimate day. For the midguts, the appearance of the E20MO coincided with the transition from larval to pupal tissue. No activity was detected in larval midguts. 3-Epiecdysone formation, however, was mainly found in larval midguts, with only marginal activity detectable in pupal midguts.     

Neurological influences on pheromone release and calling behaviour in the gypsy moth, Lymantria dispar

ABSTRACT. Surgical removal of the brain or disconnection of the last abdominal ganglion from the ventral nerve cord prevented sex pheromone release in female Lymantria dispar (L.) (Lymantriidae), as assayed by the male wing-fanning response. The calling behaviour continued to occur in individuals whose terminal abdominal ganglion had been thus isolated, however, indicating that the neural mechanisms controlling calling function independently in the last abdominal ganglion.     

Sexual dimorphism in neuronal projections from the antennae of silk moths (Bombyx mori,Antheraea polyphemus) and the gypsy moth (Lymantria dispar)

Summary The antennal lobe of both sexes of the silk moth Bombyx mori contains 55–60 ventrally located antennal glomeruli; in addition, that of the male contains a dorsal macroglomerular complex (MGC). A group of identifiable glomeruli consisting of two lateral large glomeruli (LLG) and four medial small glomeruli (MSG) is present in both sexes, but the LLG are greatly enlarged in the female. A MGC is also present in the male gypsy moth Lymantria dispar and male giant silk moth Antheraea polyphemus. The MGC in all of these species is organized into 3–4 distinct levels of glomeruli. Antennal sensory fibers were stained by cobalt backfills in B. mori, A. polyphemus, and L. dispar. Most fibers stained from cut long hairs (sensilla trichodea) projected to MGC in males and LLG in both sexes of B. mori. The distribution of fibers in the MGC of B. mori was topographically biased in that a majority of fibers from anterior branches projected medially in MGC while most fibers from posterior branches projected laterally or anteriorly. Terminal arborizations of single fibers were each restricted to a single glomerular level of the MGC. Fibers projecting to the posterior antennal center were frequently stained in cut-hair and control preparations, apparently by uptake of cobalt through intact sensilla on flagellar branches.     

Heartbeat rate modulation mediated by the ventral nerve cord in the honey bee,Apis mellifera

Summary Using impedance conversion as a recording technique, heartbeat patterns were established for intact adult honey bee workers during rest and during nectar feeding and locomotion, activities that both result in accelerating heartbeat. Heartbeat patterns were then described separately for bees in which assumed neural regulation was disrupted (the ventral nerve cord was transected), and for bees in which assumed neurohormonal regulation was disrupted (blood circulation to the abdomen was blocked). Heartbeat patterns from these latter two groups were compared with the baseline data from the former three groups. Results showed that normal modulatory patterns occurring during feeding and locomotion persisted when the neurohormonal pathway was disrupted, but the same modulation was absent when the neural pathway was disrupted. These results offer clear support for the conclusion that neural mechanisms provide a primary form of regulation of heartbeat in the honey bee.     

Effects of maternal nutrition and egg provisioning on parameters of larval hatch,survival and dispersal in the gypsy moth,Lymantria dispar L.

North American gypsy moths disperse as newly hatched larvae on wind currents in a behavior called ballooning. Because ballooning occurs before neonates begin to feed, resources used in dispersal are limited to those carried over from the egg. We show that nutritional experience of the maternal parent can influence the tendency of offspring to disperse, and that resource provisioning of eggs by the maternal parent affects the duration of the window for disperal. Offspring of females from defoliated sites had a lower tendency to balloon in a wind tunnel than larvae from females which had not experienced nutritional stress associated with host defoliation. The number of eggs in an egg mass, a reflection of the maternal parent's nutritional experience, also contributed to the predictive model for dispersal that included defoliation level. Egg weight and the levels of two yolk proteins, vitellin (Vt) and glycine-rich protein (GRP), however, had no influence of the proportion of ballooning larvae. The length of survival without food, and thus the maximum period of time for dispersal, was correlated with levels of Vt and GRP, but not with egg weight. The level of defoliation at the site from which the maternal parent was collected was not related to the longevity of offspring, nor did it have a significant effect on the levels of Vt, GRP or egg weight. Levels of hemolymph proteins arylphorin and vitellogenin in the maternal parent during the prepupal stage had no influence on levels of yolk proteins, larval longevity, or tendency to balloon.