Colour choice behaviour in the pollen beetle Meligethes aeneus (Coleoptera: Nitidulidae)

The pollen beetle Meligethes aeneus Fabricius (Coleoptera, Nitidulidae), a pest of oilseed rape (Brassica napus), is known to respond to coloured stimuli; however, current understanding of the underlying mechanisms of colour choice in this species is limited. In the present study, physiological and behavioural experiments are conducted to determine the response of the pollen beetle to colours in the field. Spectral sensitivity is measured in 10 animals using the electroretinogram technique. Light flashes (100 ms) at varied wavelengths (340–650 nm, 10‐nm steps) and at different light intensities are applied to the eye after dark adaptation. In behavioural experiments in the field, 100 water traps of varying colours (from yellow to green to blue with varying amounts of white and black added, and with known spectral reflectance) are set out on a bare soil field in May 2008. The mean spectral sensitivity curve of M. aeneus peaks at 520 nm; however, a model template fitted to the long wavelength tail of the observed curve reveals a peak at approximately 540 nm (green). A secondary sensitivity peak is observed in the ultraviolet (UV) range (370 nm). A total of 2482 pollen beetles are captured in the coloured traps. The results show that the pollen beetles' preference for yellow over other colours can be modelled as a colour opponent mechanism (green versus blue); however, further experiments are needed to specify responses to colours with higher UV reflectance. These findings may be used to optimize trap colours for monitoring to help develop integrated pest management strategies for pollen beetle control.     

Supercooling ability and cold hardiness of the pollen beetle Meligethes aeneus

Supercooling point (SCP) and cold‐hardiness of the pollen beetle Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae) were investigated. Mature eggs from the oviduct were supercooled on average to ?28.0 °C and from oilseed rape buds to ?24.4 °C; first instars were supercooled to ?21.0 °C and second instars to ?16.8 °C. Despite their high supercooling ability, none of the eggs survived 24 h exposure to ?2.5 °C. The supercooling ability of adults varied significantly among feeding and non‐feeding beetles: high SCPs prevailed during the whole warm period, being about ?12 °C; low values of SCP of ?20 °C dominated in non‐feeding beetles. In spring and autumn, beetles displayed the same acclimation efficiency: after 1 week of exposure at 2.0 °C with no access to food their SCPs were depressed equally by about 3 °C. Meligethes aeneus beetles have a different response to low temperatures depending on the season. The lowest tolerance was found in reproductively active beetles after emergence from overwintering sites; the time needed to kill 50% of individuals (Ltime₅₀) was 56.2 h at ?7 °C and the lower lethal temperature needed to kill 50% (Ltemp₅₀) after 24 h exposure was ?8.6 °C. Cold hardiness increased from midsummer to midwinter; Ltime₅₀ was 80 h in August, 182.8 h in September, and 418.1 h in January. Lethal temperature after 24 h exposure was ?9.1 °C in August and ?9.8 °C in September. In February, after diapause, the beetles started to loose their cold tolerance, and Ltemp₅₀ was slightly increased to ?9.5 °C. Hibernating beetles tolerated long exposure at ?7 °C well, but mortality was high after short exposure if the temperature dropped below ?9 °C for 24 h. Despite the season, the beetles died at temperatures well above their mean SCP; consequently, SCP is not a suitable index for cold hardiness of M. aeneus.     

Species-specific primers for predation studies of the pollen beetle, Meligethes aeneus (Coleoptera, Nitidulidae)

Species‐specific primers were developed for the pollen beetle (a pest in oilseed rape) for studies of predation by natural insect predators. Two forward and three reverse primers were designed within the mitochondrial COI gene and used in combination to amplify fragments in the size range of 163–290 bp. Remains of pollen beetle DNA were consistently detected in Pardosa spiders up to 24 h after ingestion but dropped drastically at 48 h. These primers will facilitate studies on biological control of this oilseed rape pest. Detection time was not correlated with fragment length as might be expected as the DNA gradually degrades into progressively shorter fragments over time.     

Migration to and dispersal from oilseed rape by the pollen beetle,Meligethes aeneus,in relation to wind direction

1 Host‐plant‐odour‐induced upwind anemotaxis is accepted as the main mechanism by which herbivorous insects find their host plant within an heterogenous environment, but field data supporting this hypothesis are scarce. 2 The flight directions of the pollen beetle Meligethes aeneus to and from a plot of winter oilseed rape and the direction of the wind were recorded concurrently. The beetles were sampled by eight double‐sided window traps encircling the plot. Distal sides of the traps, relative to the plot, sampled the beetles as they flew towards the plot, whereas the proximal sides of the traps sampled them as they flew away from the plot. Paired data on daily catch of beetles in distal or proximal traps and the volume of air impacting each trap were compared. 3 Correlations between daily catch of M. aeneus into distal traps and trap wind volume were negative, indicating that flights by both overwintered‐ and new‐generation insects towards the plot were upwind. 4 Correlations between daily catch of M. aeneus into proximal traps and trap wind volume varied with generation. Catch of overwintered‐generation M. aeneus was negatively correlated with trap wind volume, indicating that flight was upwind. Proximal trap catch of the new‐generation M. aeneus was weakly/modestly positively correlated with trap wind volume, indicating that flights were downwind or crosswind. 5 Understanding the effect of wind direction on flight of M. aeneus holds potential for improving the forecasting of their arrival and spatial distribution on the crop for integrated pest management.     

Response of the pollen beetle Meligethes aeneus to volatiles emitted by intact plants and conspecifics

The response of the pollen beetle Meligethes aeneus Fab. (Coleoptera, Nitidulidae) to the volatiles of undamaged plants and conspecifics was tested in a Y-tube-olfactometer-bioassay. Beetles that had hibernated preferred significantly the volatiles emitted by their most important host plant, oilseed-rape (Brassica napus L. ‘Lorar’) in the early bud-stage. However, the odour emitted by rye (Secale cereale L.), tomato plants (Solanum lycopersicum L.), and yarrow (Achillea millefolium L.) were also attractive when tested against the corresponding growing-medium. Dock plants (Rumex obtusifolius L.) and touch-me-not (Impatiens parviflora L.) possessed no attractive properties. When tested against each other, oilseed-rape was preferred significantly by M. aeneus above all other plants, with the exception of tomato. The results indicate that M. aeneus is able to locate its host plant by olfactory stimuli in the early bud-stage, i.e. in that stage in which the infestation begins in the field and when the typical yellow colour and floral scent of oilseed-rape are absent. Female pollen beetles avoided significantly the volatiles emitted by female conspecifics and an ether extract of conspecifics of mixed sex, while volatiles from males had no significant effect on females. Furthermore, males showed no preferences when responding to conspecific odour in the olfactometer. These results suggest the existence of an epideictic pheromone for M. aeneus.     

The effects of non-host plant essential oil volatiles on the behaviour of the pollen beetle Meligethes aeneus

The use of semiochemicals for manipulation of the pollen beetle Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae) is being investigated for potential incorporation into a push‐pull control strategy for this pest, which damages oilseed rape, Brassica napus L. (Brassicaceae), throughout Europe. The response of M. aeneus to non‐host plant volatiles was investigated in laboratory assays to establish whether they have any effect on host plant location behaviour. Two approaches were used. First a novel, moving‐air bioassay using air funnels was developed to compare the response of M. aeneus to several non‐host plant essential oils. The beetles avoided the host plant flowers in the presence of non‐host volatiles, suggesting that M. aeneus uses olfactory cues in host location and/or acceptance. The results were expressed as ‘repellency values’ in order to compare the effects of the different oils tested. Lavender (Lavendula angustifolia Miller) (Lamiaceae) essential oil gave the highest repellency value. In addition, a four‐arm olfactometer was used to investigate olfactory responses, as this technique eliminated the influence of host plant visual and contact cues. The attraction to host plant volatiles was reduced by the addition of non‐host plant volatiles, but in addition to masking the host plant volatiles, the non‐host volatiles were avoided when these were presented alone. This is encouraging for the potential use of non‐host plants within a push‐pull strategy to reduce the pest colonisation of crops. Further testing in more realistic semi‐field and field trials is underway.     

Effect of physiological status on olfactory and visual responses of female Hypothenemus hampei during host plant colonization

The response of Hypothenemus hampei (Ferr.) (Coleoptera, Scolytidae) (coffee berry borer, CBB) females to olfactory stimuli was demonstrated to be related with the physiological status; especially with age, whether they were mated or not, and whether they have already oviposited. A special behaviour, the abandonment of the native berry, was also found to play a major role. In contrast, response to visual stimuli (false coffee berries) was found to be independent of the physiological status of CBB females. Variations in behavioural responses to visual and olfactory stimuli are discussed in relation to colonization strategy.     

Relevance of visual and olfactory cues for host location in the mustard leaf beetle Phaedon cochleariae

The relevance of visual and olfactory cues for host‐plant location is investigated in males and females of the oligophagous mustard leaf beetle Phaedon cochleariae Fabricius (Coleoptera: Chrysomelidae). Different objects are offered in a walking arena and the behaviour of beetles is observed. Beetles orient toward vertically or horizontally striped black and white pattern independent of stripe orientation. The results suggest that contrast facilitates orientation in the field, whereas the pattern itself may be less important for host location in dense vegetation. The response to green and yellow objects is tested to investigate discrimination abilities between young (green) and mature (yellow) leaves. Beetles prefer green over yellow independent of material (cardboard or leaves of Nasturtium officinale R. Br., Brassicaceae). Preference behaviour tested in a dual‐choice contact assay coincides with visual preferences, where adults prefer young, more nutritious leaves for feeding and oviposition. Furthermore, females discriminate between visual cues of green leaves and green cardboard, whereas males do not, indicating that females are more sensitive in colour discrimination. Differences in colour wavelength influence the choice of beetle behaviour more strongly than differences in intensity. Both sexes of P. cochleariae prefer volatiles of the host plant N. officinale, whereas only females respond to the main volatile compound 2‐phenylethyl isothiocyanate. Given a choice between visual and olfactory cues, males orientate towards the colour cues, whereas females do not show any preferences. In males, visual cues may thus override olfactory cues, whereas, in females, both are equally important, which may reflect different ecological requirements and/or physiological abilities.     

Electrophysiological and behavioural responses of the pollen beetle, Meligethes aeneus, to volatiles from a non-host plant, lavender, Lavandula angustifolia (Lamiaceae)

A semiochemical based push-pull strategy for control of oilseed rape pests is being developed at Rothamsted Research. This strategy uses insect and plant derived semiochemicals to manipulate pests and their natural enemies. An important element within this strategy is an understanding of the importance of non-host plant cues for pest insects and how such signals could be used to manipulate their behaviour. Previous studies using a range of non-host plants have shown that, for the pollen beetle Meligethes aeneus (Coleoptera: Nitidulidae), the essential oil of lavender, Lavandula angustifolia (Lamiaceae), was the most repellent. The aim of this study was to identify the active components in L. angustifolia oil, and to investigate the behaviour of M. aeneus to these chemicals, to establish the most effective use of repellent stimuli to disrupt colonisation of oilseed rape crops. Coupled gas chromatography-electroantennography (GC-EAG) and gas chromatography-mass spectrometry (GC-MS) resulted in the identification of seven active compounds which were tested for behavioural activity using a 4-way olfactometer. Repellent responses were observed with (±)-linalool and (±)-linalyl acetate. The use of these chemicals within a push-pull pest control strategy is discussed. Handling Editor: Yvan Rahbe.     

Laboratory and field response of the pollen beetle, Meligethes aeneus, to the odour of oilseed rape

Abstract. The behavioural response of adults of the pollen beetle, Meligethes aeneus F. (Coleoptera: Nitidulidae), to the odour of one of their host-plants, oilseed rape ( Brassica napus L.), was observed in the laboratory using a fourarmed airflow olfactometer, and under semi-natural conditions using a markrelease–recapture technique. Oilseed rape leaf and flower odour was attractive to pollen beetles in the olfactometer and they dispersed upwind from the centre of a circular array (radius 20 m) of yellow water traps, baited with extracts of oilseed rape leaves or flowers. The dispersal of pollen beetles in the absence of oilseed rape odour was predominantly downwind and crosswind. The percentage recapture of pollen beetles was significantly greater in traps baited with rape flower extracts than leaf extracts. Pollen beetles can use odour-mediated upwind anemotaxis to locate oilseed rape plants from distances of at least 20 m over open ground.     

视觉和嗅觉信号对果蝇食物搜寻行为的协同作用

为了探索视觉和嗅觉信号在昆虫食物搜寻过程中的作用, 本研究利用杨梅和橘子为引诱物, 在实验室条件下测定了嗅觉和视觉信号诱集到的黑腹果蝇Drosophila melanogaster数量, 分析了嗅觉经历对果蝇嗅觉和视觉食物搜寻的影响。发现同源性嗅觉和视觉信号存在的杨梅诱集到的果蝇数量显著大于单一的视觉信号和嗅觉信号, 但异源性嗅觉和视觉信号组合诱集到的果蝇数量和单独的嗅觉信号相似。嗅觉信号预处理不仅能够显著增加嗅觉信号诱集到的果蝇数量, 其中杨梅嗅觉信号对杨梅预处理果蝇的吸引能力与视觉和嗅觉信号存在的杨梅相似, 而且异源性嗅觉和视觉信号组合诱集到的预处理果蝇数量也不低于视觉和嗅觉信号存在的杨梅。另外杨梅嗅觉信号预处理也能够显著增强杨梅视觉信号诱集到的果蝇数量。但嗅觉预处理并不会改变同源性视觉和嗅觉信号组合诱集到的果蝇数量。本研究表明, 果蝇同时利用视觉和嗅觉信号进行食物搜寻, 因此同源性视觉和嗅觉信号在果蝇诱集过程中具有协同作用。另外果蝇具有较强的记忆和学习能力, 能够将记忆中的嗅觉信号应用于食物搜寻。本研究结果不仅有利于我们了解果蝇在自然状态下的食物搜寻机制, 而且有利于开发更有效的果蝇新型诱捕器。     

The roles of olfaction and vision in host-plant finding by the diamondback moth, Plutella xylostella

Abstract.  The relative roles of olfaction and vision in the crepuscular host-finding process of a major lepidopteran pest of cruciferous crops, the diamondback moth Plutella xylostella are investigated in a series of laboratory and semi‐field experiments. Flying female moths use volatile plant chemical cues to locate and to promote landing on their host, even in complex mixed-crop environments in large cages. Multiple regression analysis shows that both the plant position (front, middle or back rows) and the type of plant (host plant, nonhost plant) are needed to explain the distribution of insects in such a mixed-crop situation. This strong plant position effect indicates that, when host plants are present in a mixture, foraging P. xylostella are more likely to alight on the first row of the plants. The findings are discussed with regard to current theories of host-plant location by phytophagous insects and the possible implications for integrated pest management.     

Behavioural responses in three ichneumonid pollen beetle parasitoids to volatiles emitted from different phenological stages of oilseed rape

Pollen beetles (Meligethes spp.; Coleoptera: Nitiduliae) are a major pest of oilseed rape, Brassica napus L. (Brassicaceae) in northern Europe. Phradis interstitialis Thomson, P. morionellus Holmgr., and Tersilochus heterocerus Thomson (Hymenoptera: Ichneumonidae) are among the most frequent pollen beetle parasitoids. These three species differ in temporal occurrence, as well as in preferred host stage. The behavioural responses of female parasitoids to odours from oilseed rape at bud and flowering stage were evaluated in two‐choice experiments. The role of visual stimuli was examined by combining green and yellow colours with odour stimuli. All three species were attracted to odours from the bud stage of oilseed rape. Tersilochus heterocerus was attracted to odours of flowering rape, but the two Phradis species avoided the flower odours. However, when the odours of flowering rape were combined with yellow, and odours of the bud stage were combined with green, P. interstitialis was equally attracted to both stimuli, and T. heterocerus showed an increased preference for flower odours, while no effect of colours could be found in P. morionellus. The observed differences in responses between the parasitoids may reflect differences in their biology and may be involved in the niche segregation of these often coexisting species. The volatile blends released from the two phenological stages were identified and compared. Clearly, odours can be reliable cues for differentiating between oilseed rape in the bud and flowering stage. Of 20 identified compounds, 18 were released at a significantly higher rate from flowering plants. The terpenes sabinene, myrcene, limonene, and (E,E)‐α‐farnesene were the dominant volatiles in the bud and flower headspace. A group of aromatic compounds including benzaldehyde, methyl benzoate, and phenyl acetaldehyde were mainly released from flowering rape.     

Visual and olfactory disruption of orientation by the western pine beetle to attractant-baited traps

Olfactory deterrents have been proposed as tree protectants against attack by bark beetles, but their development has been hindered by a lack of knowledge of host selection behavior. Among the primary tree-killing (aggressive) Dendroctonus, vision appears to be an integral part of the host selection process. We evaluated the importance of vision in host finding by D. brevicomis LeConte, and our ability to affect it by modifying the visual stimulus provided by attractant-baited multiple-funnel traps. White-painted traps caught 42% fewer D. brevicomis than black traps in California, USA (P < 0.05). Visual treatments were less effective (P < 0.0001) than olfactory disruptants (verbenone with ipsdienol), which reduced catch by about 78%. When combined, olfactory and visual disruptants resulted in 89% fewer D. brevicomis being caught, but this combination was not more effective than olfactory disruptants alone (P > 0.05). Our results demonstrate that the visual component of D. brevicomis host finding behavior can be manipulated, but that D. brevicomis may be more affected by olfactory than visual disruptants. In contrast, visual disruption is more pronounced in the southern pine beetle, Dendroctonus frontalis Zimmermann, suggesting that non-insecticidal tree protection strategies for these related species should differ.     

Influence of visual and olfactory cues on field trapping of the pollen beetle, Astylus atromaculatus (Col.: Melyridae)

Field trapping experiments investigated the response of the pollen beetle Astylus atromaculatus to visual and olfactory cues during a 3-year period, 1999–2001. The visual preference of the pollen beetle was determined using yellow, white, blue, green and red water traps. The yellow trap was most attractive, capturing 56% of the total beetles trapped, with 30% caught by the blue and white traps, while 14% was caught by the red and green traps. The response of the beetle to olfactory cues was then evaluated by using the yellow water trap with three antennally active components identified in the volatiles of sorghum panicles by coupled gas chromatography (GC)–electroantennographic detection and GC–mass spectrometry. These components were 2-phenylethanol, benzyl alcohol and linalool. There were no significant colour × chemical compound interactions and traps baited with 2-phenylethanol captured significantly more beetles than unbaited traps, irrespective of trap colour, demonstrating the effectiveness of olfactory cues in trapping the pollen beetle. Traps baited with 2-phenylethanol were more attractive than and caught more beetles than traps baited with linalool. 2-Phenylethanol had the greatest effect on the relatively unattractive blue trap, confirming the importance of olfactory cues mediating A. atromaculatus attraction .     

Orientation responses of the grasshopper, Melanoplus sanguinipes, to visual, olfactory and wind stimuli and their combinations

Prereproductive adults of the grasshopper, Melanoplus sanguinipes (F.) (Orthoptera, Acrididae), demonstrated orientation and movement towards both visual and olfactory stimulus sources in a still-air chamber. Visual stimuli (wheat and lima bean foliage, vertical black or yellow-green stripes, and a yellow-green broad leaf pattern) were approached more frequently than the control white background surface. Olfactory stimuli (chopped wheat foliage and a four-component, synthetic, grass odor blend of volatiles) elicited an even greater positive response than the visual stimuli. Changing the proportions of the four volatiles in the blend significantly reduced positive orientation responses to the stimulus source. Visual cues of wheat foliage and olfactory cues of either chopped wheat odor or the grass odor blend gave greater responses when combined than when presented separately.In flowing air or wind, nearly all insects demonstrated a rapid positive response to odors of chopped wheat and the grass odor blend, significantly greater than the response to the same stimuli in still air. However, positive responses to visual cues were not significantly greater in wind than in still air. When combined with the olfactory stimuli in flowing air, visual cues did not increase the incidence of response. Grasshoppers responding to grass odors in wind moved more rapidly and directly toward the source, and stopped less often and for shorter durations than insects responding to odor in still air or to visual cues.We conclude from these studies that M. sanguinipes adults show orientation behavior to both visual and olfactory stimuli from food plant sources, although leaf odors elicit a stronger positive response particularly when carried by wind.     

Role of Olfactory and Visual Cues in the Attraction/Repulsion Responses to Conspecifics by Gregarious and Solitarious Desert Locusts

Desert locusts [Schistocerca gregaria Forskål (Orthoptera, Acrididae)] change phase in response to population density: solitarious insects avoid one another, but when crowded they change to the gregarious phase and aggregate. The attraction/repulsion responses of gregarious and solitarious locusts maintain phase differences in locust populations. Despite considerable research, the cues for aggregation are poorly understood; moreover, the repulsion response of solitarious locusts has not previously been investigated. This study analyzes the role of visual and olfactory stimuli in triggering these different responses to conspecifics. Isolation-reared insects were repelled by both olfactory and visual stimuli from other locusts. Crowd-reared insects were attracted by the combination of olfactory and visual cues. In addition, olfactory stimuli affected other behaviors in both phases, and behavioral differences between isolation- and crowd-reared locusts were clear even in the absence of conspecifics. The sensory and neurological mechanisms underlying these responses are not well understood and will form the basis for neurobiological investigations of locust phase.     

Behavioral responses of the small hive beetle,Aethina tumida,to odors of three meliponine bee species and honey bees,Apis mellifera scutellata

Recent studies have shown that honey bees, bumble bees, and some meliponine bee species of the genera Trigona, Meliponula, and Dactylurina are hosts of the small hive beetle (SHB) Aethina tumidaMurray (Coleoptera: Nitidulidae), a pest of honey bee colonies in various regions of the world. Olfaction has been implicated in SHB infestations of honey bee and bumble bee colonies. We used olfactometer bioassays to investigate responses of adult male and female SHBs to odors from intact colonies and separate hive components (pot honey, pot pollen, cerumen, and propolis) of three African meliponine bee species, Meliponula ferruginea (Lepeletier) (black morphospecies), M. ferruginea (reddish brown morphospecies), and Meliponula bocandei (Spinola) (Hymenoptera: Apidae). Although both sexes of the beetle strongly preferred intact colony, pot honey, and pot pollen odors, there was no evidence of attraction to propolis and cerumen odors from the three meliponine bee species. Both sexes of SHB also strongly preferred odors from honey bees, Apis mellifera L. (Hymenoptera: Apidae), over odors from the three meliponine bee species. Our results provide substantial evidence of the host potential of African meliponine bees for the SHB, and we discuss this complex association of the SHB with species within the Apidae family.     

Sequestration and metabolism of host-plant flavonoids by the Pale Grass Blue, Pseudozizeeria maha (Lepidoptera: Lycaenidae)

Flavonoids in the eggs, larvae, pupae and adult butterflies of Pseudozizeeria maha were identified and quantified. Isovitexin (apigenin 6- C -glucoside) was the dominant flavonoid detected in pupae and imagines, whereas saponarin (isovitexin 7- O -glucoside) was the dominant flavonoid in larvae. Saponarin, biotransformed from isovitexin by the larvae, was found to be reconverted into isovitexin at the pupal stages. The total amounts of accumulated flavonoids differed between sexes, particularly in adults where the bodies of females had a much higher flavonoid concentration than that of the males. These flavonoids were also detected in the eggs. These results indicate the possibility that flavonoids are utilized in not only wing coloration but also as a chemical defense compound against natural threats.