The effect of photoperiod on the calling behaviour of virgin females of the true armyworm,Pseudaletia unipuncta (Haw.) (Lepidoptera: Noctuidae)

The effect of photoperiod on the calling behaviour of Pseudaletia unipuncta virgin females was examined under five different photoperiodic regimes at 25°C, 65% r.h. The age at which females called for the first time following emergence varied with photoperiod; generally calling was later under long-scotophase conditions. However under a 6 h scotophase there was also a delay in calling and >63% of the females tested never called. There was a considerable variability in the daily calling patterns between the different photoperiods, and the mean onset time of calling was not constant with either “lights on” or “light off”. There was however a constancy of the mean onset time of calling relative to the mid-point of both the photo- and scotophase at all photoperiods tested, indicating that females could measure the absolute duration of either the photo- or scotophase. Transferring females from one photoperiodic condition to another once calling had been initiated, determined that it was the “lights off” signal that P. unipuncta females used to phase set the clock governing circadian calling behaviour.Females subjected to a decrease of 4 or 6 h in the length of the scotophase following the initiation of calling required several days to adjust to the new photoperiodic regime and a high proportion of females did not call during the night following the transfer. A 4 or 6 h increase in the scotophase did not inhibit calling on the night following the transfer but females still required several days to adjust completely. However, females experiencing a 2 h increase or decrease in the duration of the scotophase were able to maintain normal calling behaviour. The results of these experiments are discussed in relation to the seasonal biology of the true armyworm and the hypothesis that this is a migrant species.     

Primary and continuous midgut cell cultures from Pseudaletia unipuncta (Lepidoptera: Noctuidae)

Midgut epithelial cells were isolated from fifth-instar Pseudaletia unipuncta larvae by collagenase treatment of midgut tissue, and cultured in TNM-FH medium. Long-term continuous culture and maintenance of midgut cells were achieved with P. unipuncta armyworm intestinal cells. Several cells lines were obtained from these P. unipuncta primary cultures, and they have been subcultured and maintained for over 24 mo. The three major midgut cell types were present in the cultures, including stem (regenerative), columnar, and goblet cells. In vitro morphogenesis and differentiation of columnar and goblet cells from stem cells were observed. There appeared to be a cycle of cell death of goblet and columnar cells followed by their replacement from stem cells every 7-8 wk. After approximately six passages, the cell density in T-flasks appeared to be somewhat constant, reaching 10(3)-10(4) cells per milliliter of medium. The columnar cells are round to rectangular in shape and possess a brush border, while the goblet cells have a classic flask-like shape with a central cavity. Peritrophic membrane-like secretions were observed in all the culture flasks. Infection of these cells with multiply embedded nucleopolyhedrovirus was confirmed, and we conclude that these midgut cells can be used as an in vitro model system to study early events in baculovirus infection.     

Biological activity of essential oils from seven Azorean plants against Pseudaletia unipuncta (Lepidoptera: Noctuidae)

Ten essential oils from seven Azorean plant species were evaluated for their insecticidal, ovicidal, feeding‐deterrence and growth inhibition activities against Pseudaletia unipuncta. The oils of Laurus azorica (leaves), and Juniperus brevifolia (leaves) showed strong moderate insecticidal effect on fourth‐instar larvae causing 93.3% and 46.7% mortality, respectively. Juniperus brevifolia (leaves), L. azorica (leaves), Persea indica (leaves), Hedychium gardnerianum (leaves) and Pittosporum undulatum (fruits and leaves) significantly affected the hatching of P. unipuncta eggs (<8% eclosions). Five oils showed significant feeding deterrent activities (L. azorica, 92.4%, J. brevifolia, 93.6%, P. undulatum leaves, 95.5% and fruits, 83.8% and H. gardnerianum, 88.2%). All of the essential oils tested, significantly inhibited the larval growth after 5 days of feeding on the treated diet. Essential oils from L. azorica and J. brevifolia were the most potent growth inhibitors among the oils tested, producing a decrease in the initial larva weight (?14.8 and ?14.5 mg, respectively). Our results indicate that L. azorica (leaves), J. brevifolia (leaves), P. indica (leaves), H. gardnerianum (leaves), and P. undulatum (leaves and fruits) can be exploited for the development of bioactive compounds as a new source of agrochemicals. Further emphasis on isolation and identification of active constituents can be useful to develop new environment‐friendly insect control agents.     

Susceptibility to insecticides of Glyptapanteles militaris (Hymenoptera: Braconidae), a Parasitoid of Pseudaletia unipuncta (Lepidoptera: Noctuidae)

The susceptibility of cocoons and adults of Glyptapanteles militaris (Walsh) were studied. One organophosphate insecticide (trichlorfon), one organochlorine insecticide (endosulfan), one pyrethroid (deltamethrin) and a commercial formulation of Bacillus thuringiensis subsp. Kurstaki were selected for testing. All the tests were carried out with fresh solutions of commercial insecticides applied on host larvae at the recommended concentration. One- and 6-day-old cocoons were sprayed with the insecticide solutions by means of a Potter Tower and held for adult emergence. Adults were exposed to residues of insecticides inside plastic vials. The B. thuringiensis formulation had no harmful effect on the cocoons nor on the adults. Trichlorfon and endosulfan were highly toxic to all the tested stages. Deltamethrin was highly toxic to adults but relatively safe when applied on parasitoid cocoons. Based on these results, field applications of deltamethrin would be least disruptive of tested insecticides to populations of G. militaris.     

Serological study on the transmission of a granulosis virus of the armyworm,Pseudaletia unipuncta (Lepidoptera: Noctuidae), to other lepidopterous species

The synergistic (Hawaiian) strain of a granulosis virus (GV) from the armyworm, Pseudaletia unipuncta, was transmitted to Spodoptera exigua, Autographa californica, and Trichoplusia ni. The viruses isolated from these hosts were tested by radial double-immunodiffusion (RDD) and immunoelectrophoresis (IE) for their relationship to the original virus. Untreated and heat-treated virus inclusion bodies (capsules) were compared for their antigenic properties but no differences were detected. The antiserum elicited against the whole capsule was more sharply specific for the antigenic determinants than the one elicited against the dissolved capsule proteins. The viruses obtained from S. exigua and T. ni elicited precipitin lines that differed from those of the P. unipuncta GV in their electrophoretic mobility with the one-trough IE method; however, with the two-trough method, the lines fused indicating that the antigens were identical. The major precipitin line indicating identity of the viruses wasthat produced by the synergistic factor (SF) purified from the capsule proteins of the synergistic GV strain. The presence of SF in the GV produced in alternate hosts indicated that its production was virus directed. The SF was not detected in the GVs of Laspeyresia pomonella and Pieris rapae and in the nonsynergistic Oregonian GV of P. unipuncta. A field-collected GV from S. exigua exhibited a different precipitin pattern from that of the synergistic GV, but one of the precipitin lines shared partial relationship to the SF.     

Susceptibility of Pseudaletia unipuncta (Lepidoptera: Noctuidae) to entomopathogenic nematodes (Rhabditida: Steinernematidae and Heterorhabditidae) isolated in the Azores: effect of nematode strain and host age

The armyworm, Pseudaletia unipuncta (Haworth), is a serious pest to the Azores's pastures. In laboratory bioassays we tested the susceptibility of this insect to entomopathogenic nematodes isolated in Azores: Steinernema carpocapsae Az20, Az150, and A48 strains, S. glaseri Az26 strain and Heterorhabditis bacteriophora Az33 strain. The A48, Az20, and Az150 strains caused parasitism rates of 96.6, 90, and 53.3%, and mortality rates of 63.3, 46.6, and 23.3%, respectively, to sixth instar. The Az33 strain caused a parasitism rate of 73.3% and a mortality rate of 40%; whereas, the Az26 strain caused a parasitism rate of 40% and no mortality. A linear response dose-parasitism with a positive regression (r2 = 0.993) was observed in insects exposed to S. carpocapsae Az150 strain. Positive regressions were also observed between mortality and dose rate for S. carpocapsae A48 (r2 = 0.980), Az20 (r2 = 0.956), and Az150 (r2 = 0.963) strains, and H. bacteriophora Az33 strain (r2 = 0.999). Fourth instars were the most susceptible to the A48 strain, followed by the fifth instars, while the sixth instars were the less susceptible, with LD50 values of 26.2, 62.8, and 320.7 infective juveniles, respectively. The lethal time for each of the tested instars was 32.3, 35.5, and 49.2 h, respectively. The invasion rate was 33.5, 28.2, and 40.8 nematodes per treated larvae in the fourth, fifth, and sixth instars, respectively.     

Differences between the pathogenic processes induced by Steinernema and Heterorhabditis (Nemata: Rhabditida) in Pseudaletia unipuncta (Insecta: Lepidoptera)

Larvae of Pseudaletia unipuncta are moderately susceptible to infections caused by entomopathogenic nematodes, being a desirable host to study pathogenic processes caused by Heterorhabditis bacteriophora, Steinernema carpocapsae, and Steinernema glaseri and their associated bacteria. The ability of the infective stage of these nematodes to invade hosts is quite different. S. carpocapsae invades the highest number of insects and presents the highest penetration rate, followed by H. bacteriophora. Regression analysis between the number of insects parasitized and the number of IJs counted per insect, over time, showed a high correlation for S. carpocapsae whereas for H. bacteriophora it was low. Dose-response was most evident at a concentration below 100 IJs per insect on H. bacteriophora, whereas on S. carpocapsae it was found for doses ranging from 100 to 2,000 IJs. Student's t test analysis of dose-response showed parallel, yet unequal, slopes for both strains of H. bacteriophora, whereas distinct regressions were obtained for S. carpocapsae and S. glaseri, thus, evidencing each species develop a distinct pathogenic process. Insects injected with Photorhabdus luminescens died within 50 h after injection, whereas those treated with X. nematophila died much later. Moreover, the mortality in insects exposed to H. bacteriophora complex and injected with P. luminescens was close, but insects injected with bacteria died faster. Insect mortality in treatments with complexes S. carpocapsae and S. glaseri was significantly higher than that which was observed in insects injected with symbiotic bacteria.     

Pupal Period Affects Calling Behavior of the Wheat Moth, Pseudaletia sequax (Lepidoptera: Noctuidae)

The effects of pupal period and age on calling behavior of virgin females of Pseudaletia sequax were determined. Calling behavior of groups of females of similar age of pupation was observed every 10 min for six calling days. Females were considered calling when they assumed a characteristic position: wings and abdomen elevated, displaying the ovipositor and presumably releasing pheromone. Calling behavior was discontinuous. Females with a shorter pupal period took more time to start calling than females with a longer pupal period and called for a longer period. On the first day of calling, females took more time to initiate calling, on average during the seventh hour of the scotophase, whereas on the second day of calling, most females initiated calling during the fifth hour of the scotophase. On subsequent days of calling, the average time to initiate calling changed to the fourth hour of the scotophase. The mean time of calling per day decreased significantly with the number of days since first calling, and the mean number of calling bouts increased with days since first calling.     

Sensilla on the antennal flagellum of the sawyer beetles Monochamus notatus (Drury) and Monochamus scutellatus (Say) (Coleoptera: Cerambycidae)

The antennae of the sawyer beetles Monochamus notatus and M. scutellatus were examined with the light and scanning electron microscopes to determine the types, number, distribution and innervation of the sense organs. Nine types of sensilla are described. Both short, thin-walled pegs (sensilla basiconica) and reversely curved thick-walled hairs (sensilla trichodea) are chemoreceptors. There are three types of long, thick-walled hairs (sensilla chaetica) which may be mechanoreceptors. One of these is modified in males to form unique snail-shaped pegs. A few dome-shaped organs, probably campaniform sensilla, were found. In addition to sense organs, many glands occur in association with the sensilla, and the antennae are well supplied with dermal glands connected by canals to small pores on the surface.     

Spatial representation of odours in the antennal lobe of the moth Spodoptera littoralis (Lepidoptera: Noctuidae)

Glomeruli within the antennal lobe (AL) of moths are convergence sites for a large number of olfactory receptor neurons (ORNs). The ORNs target single glomeruli. In the male-specific cluster of glomeruli, the macroglomerular complex (MGC), the input is chemotypic in that each glomerulus of the MGC receives information about a specific component of the conspecific female sex pheromone. Little is known about how neurons that detect other odorants arborize in and amongst glomeruli. The present study focuses on how sex pheromones and biologically relevant semiochemicals are represented in the ALs of both sexes of the moth Spodoptera littoralis. To assess this, we optically measured odour-evoked changes of calcium concentration in the ALs. Foci of calcium increase corresponded in size and shape with anatomical glomeruli. More than one glomerulus was normally activated by a specific non-pheromonal odorant and the same glomerulus was activated by several odorants. All odorants and pheromone components tested evoked unique patterns of glomerular activity that were highly reproducible at repeated stimulations within an individual. Odour-evoked patterns were similar between individuals for a given odorant, implicating a spatial olfactory code. In addition, we demonstrated that activity patterns evoked by host-plant related volatiles are similar between males and females.     

Interference between two nuclear polyhedrosis viruses of the armyworm,Pseudaletia unipuncta [Lep.: Noctuidae]

The larva ofPseudaletia unipuncta (Haworth) is susceptible to 2 nuclear polyhedrosis viruses (NPV), the typical (TNPV) and the hypertrophy (HNPV) strains. The interference between the 2 viruses was studied primarily in the tracheal and fat tissues. When both viruses were microfed simultaneously, no larva had mixed virus infections and most were infected with TNPV. In order to obtain 50% infection by each virus the inoculum of HNPV had to be increased by 1000 polyhedra/ml at the LD₅₀ value of TNPV. Mixed infection resulted when TNPV was inoculated 1 and especially 2 days after HNPV inoculation. In mixed infection, each virus attacked certain areas of the tracheae instead of random cells. These results indicated an interference or antagonism between the 2 viruses. The possible cause for the interference is discussed.

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Résumé La larve dePseudaletia unipuncta (Haworth) est sensible à 2 virus à polyèdres nucléaires (NPV), la souche typique (TNPV) et la souche causant une hypertrophie (HNPV). L'interférence entre ces 2 virus a été étudiée essentiellement dans les trachées et le corps adipeux. Si les deux virus sont inoculés simultanément par micro ingestion aucune larve ne présente d'infection mixte et la plupart d'entre elles est infectée par le TNPV. Pour obtenir 50% d'infection par chacun des virus, l'inoculum de HNPV doit être accru de 1000 polyèdres par ml par rapport à l'inoculum provoquant la DL 50 du TNPV. Une infection mixte est enregistrée quand le TNPV est inoculé 1 jour et de préférence 2 jours après l'introduction du HNPV. Dans ce cas d'infection mixte chaque virus attaque certaines régions des trachées et non pas des cellules réparties au hasard. Ces résultats prouvent une interférence ou un antagonisme entre les 2 virus, dont la cause est discutée.

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Portion of a dissertation submitted by the senior author for the degree of doctor of philosophy from the University of California, Berkeley. Part of the study was supported by a NSF grant provided to the junior author.     

Dose-dependent response characteristics of antennal lobe neurons in the male moth Agrotis segetum (Lepidoptera: Noctuidae)

Responses of antennal lobe neurons to different amounts of the female-produced pheromone blend and to its individual components were investigated in Agrotis segetum males using intracellular recording methods. We identified three physiological types of antennal lobe neurons, categorized according to their response thresholds to single pheromone components and to the blend: generalist neurons, component-specific neurons and blend-specific neurons. Response and specificity of antennal lobe neurons were largely dose dependent. In most cases specific responses occurred only at low stimulus amounts, while increasing concentrations often resulted in an increase of the number of pheromone stimuli to which the neuron responded. Dose-response relationships often differed between different stimuli activating a neuron. Accepted: 24 May 1997     

Sense organs on the antennal flagellum of the mimosa webworm,Homadaula anisocentra (Lepidoptera,Glyphipterygidae)

Five different types of sense organs were found on the antennal flagellum of Homadaula anisocentra. These were (1) tactile hairs; (2) thick-walled chemoreceptors; (3) thin-walled chemoreceptors of several kinds; (4) styloconic chemoreceptors and (5) small chemoreceptor pegs in shallow depressions. No coeloconic sense organs were seen.     

宽胫夜蛾成虫触角和下唇须感器的超微形态

触角和下唇须是昆虫头部的重要感觉器官,其上被有多种类型的感器。宽胫夜蛾Protoschinia scutosa是一种重要的农业害虫,但其触角和下唇须感器研究较少。本研究利用扫描电子显微镜观察了宽胫夜蛾的触角和下唇须感器的类型、分布和超微形态。宽胫夜蛾的雌雄触角均为丝状,雌虫触角鞭节明显短于雄虫。雌雄触角均分布有8种类型感器,分别为Bhm氏鬃毛、鳞形感器、毛形感器、刺形感器、耳形感器、锥形感器、腔锥形感器和栓锥形感器。其中,毛形感器数量最多,且雄性毛形感器的数量明显多于雌性。宽胫夜蛾的下唇须为长管状,雌雄下唇须长度无显著差异。共观察到5种类型感器,即毛形感器、钟形感器、鳞形感器、刺形感器和棒形感器。毛形感器分布在下唇须基部第1节和端部第3节的陷窝器内,钟形感器位于第1节毛形感器簇之中,鳞形感器在各节均有分布,刺形感器和棒形感器只见于第3节。并对宽胫夜蛾的触角和下唇须感器的功能进行初步的推测和讨论。为昆虫感器功能分析、分类学和行为学研究提供依据。     

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     

Sex pheromone and plant-associated odour processing in antennal lobe interneurons of male Spodoptera littoralis (Lepidoptera: Noctuidae)

Antennal lobe interneurons of male Spodoptera littoralis (Boisd.) were investigated by using intracellular recording and staining techniques. Physiological and morphological characteristics of local interneurons and projection neurons responding to sex pheromone and plant-associated volatiles are described. The interneurons identified were divided into three groups, depending on their physiological response characteristics. Both types of interneurons, local interneurons and projection neurons, were described in all three groups. 1. Interneurons responding exclusively to sex pheromone stimuli, displayed different degrees of specificity. These neurons responded to either one, two, three or all four of the single sex pheromone or sex pheromone-like compounds tested. Most of these neurons also responded to a mixture of the two pheromone components present in the female S. littoralis blend. Two local interneurons and one projection neuron were identified as blend specialists, not responding to the single female produced sex pheromone components, but only to their mixture. Five pheromone specific projection neurons arborized in one or more subcompartments of the macroglomercular complex (MGC) and some of them had axonal branches in the calyces of the mushroom body and in different parts of the lateral protocerebrum. 2. Interneurons responding only to plant-associated volatiles varied highly in specificity. Neurons responding to only one of the stimuli, neurons responding to a variety of different odours and one neuron responding to all stimuli tested, were found. Three specialized local interneurons had arborizations only in ordinary glomeruli. One specialized and three less specialized local interneurons had arborizations within the MGC and the ordinary glomeruli. The projection neurons responding only to plant-associated volatiles had mostly uni- or multiglomerular arborizations within the ordinary glomeruli. 3. Interneurons responding to both sex pheromones and plant-associated stimuli varied in specificity. Individual interneurons that responded to few plant-associated odours mostly responded to several pheromone stimuli as well. Projection neurons responding to most of the plant-associated volatiles also responded to all pheromone stimuli. Two local interneurons responding to both stimulus groups, arborized within the MGC and the ordinary glomeruli. Projection neurons mostly arborized in only one ordinary glomerulus or in one compartment of the MGC. The variation in specificity and sensitivity of antennal lobe interneurons and structure-function correlations are discussed.     

Sense organs on the antennal flagellum of mayflies (Ephemeroptera)

A few very small sensory pegs with the characteristics of chemoreceptors are present in both males and females on the antennal flagellum of the mayflies, Stenacron interpunctatum (Say)

1 Stenacron interpunctatum (Say) is Stenonema interpunctatum (Say) of earlier literature.

and Cloeon sp. They are situated near the proximal end of the flagellum on its ventro-lateral surface and have escaped the attention of earlier investigators.