Volatiles released from foliar extract of host plant enhance landing rates of gravid Polygonia c‐album females,but do not stimulate oviposition

The role of olfactory cues for host search is much less investigated in day‐active butterflies than in their relatives, the nocturnal moths. The goal of this study was to investigate whether host‐plant volatiles from foliar extracts of hop, Humulus lupulus L. (Cannabaceae), evoke electroantennographic (EAG) responses, increase landing rates, and stimulate egg‐laying behavior of gravid Polygonia c‐album L. (Lepidoptera: Nymphalidae) females. Eighty‐nine volatile compounds were detected in a non‐concentrated methanol extract of hop by gas chromatography–mass spectrometry, 11 of which elicited an EAG response. Concentration of the crude extract significantly reduced landing rates on artificial leaves treated with the sample due to loss of volatile compounds, but after landing the oviposition response of gravid females was not affected. A mixture of eight commercially available EAG‐active volatiles increased the landing rate of gravid females to their source but did not act as oviposition stimulants. Dividing the volatile compounds into two groups – consisting of (1) hexanal, (E)‐2‐hexenal, octanal, nonanal, and decanal, and (2) sulcatone, humulene, and benzyl alcohol – obliterated effectiveness, revealing synergism between compounds. Although volatiles did not stimulate oviposition, they significantly contributed to the distribution of eggs by increasing the landing rates on treated artificial leaves.     

Flight Tunnel Responses of Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) to Olfactory and Visual Stimuli

Cohorts of mass-reared, naive Diachasmimorpha longicaudata, parasitoids of tephritid fruit flies, were released in a laminar airflow wind tunnel to study their responses to visual and olfactory stimuli associated with their host habitat. Parasitoids were five times more likely to land on yellow plastic spheres emitting the odor of ripe, guava fruit (Psidium guajava L.) than to spheres emitting clean air. The rate of landing was not modified by the presence of green artificial leaves adjacent to the spheres in the tunnel or by the inclusion of green leaf volatiles emanating with the guava odors. However, hovering activity was twice as frequent around spheres adjacent to artificial leaves than around isolated spheres. Oviposition activity on spheres emitting guava odor was not affected by the presence of artificial leaves nor by green leaf volatiles. This suggests that inexperienced D. longicaudata may be instinctively attracted to foliage and to fruit odor but that landing (arrestment) and oviposition are influenced more by odor than by the appearance of fruit or foliage. D. longicaudata are not instinctively attracted to larvae of Bactrocera dorsalis in the absence of host-habitat stimuli. More wasp activity occurred around oviposition units containing larvae if the odor of ripe/overripe guava was present. Successful wasp reproduction occurred only in units with guava odor.     

Physical and chemical factors influencing landing and oviposition by the cabbage root fly on host-plant models

Various plant models were used in both choice and no-choice tests in the laboratory to assess landing and oviposition preferences of the cabbage root fly,Delia radicum (L.). The main factor governing the site most suitable for landing was the conspicuousness of the object and not its shape. Oviposition was influenced considerably by the pre-conditioning of the females. Deprived females laid eggs even when denied access to both host plant chemicals and host-plant models. The dominant role of contact chemical stimuli in host acceptance was reconfirmed, but only a combination of physical and chemical stimuli appeared capable of eliciting normal oviposition. The combination of contact chemical stimuli and the presence of a stem on the test model had a synergistic effect on the numbers of eggs laid in both choice and no-choice situations. In choice bioassays, female cabbage root flies distinguished between models of different shapes, heights and sizes. The size and shape of the models appeared to be perceived in part after the flies had landed.     

Volatile and Contact Chemicals Released by Nezara viridula (Heteroptera:Pentatomidae) Have a Kairomonal Effect on the Egg Parasitoid Trissolcus basalis (Hymenoptera: Scelionidae)

The responses of females of the egg parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) to volatile and contact chemicals from its host Nezara viridula (L.) (Heteroptera: Pentatomidae) were investigated in a Y-tube olfactometer and under open arena conditions. In the Y-tube tests, volatiles from virgin males and from females in a preovipositional state attracted T. basalis females, while volatiles from host virgin females did not. In an open arena, traces left by N. viridula adults in different physiological conditions function as contact cues inducing the wasps to remain longer in the arena and to change the pattern of their walking behavior. However, only contact kairomones from N. viridula mated females in a preovipositional condition induced an arrestment response characterized by an increase in patch searching time and turning rates and a reduction in linear speed. The chemical ecological implications of these results on this host–parasitoid association are discussed.     

Pre- and post-landing response of the parasitoid Encarsia formosa to whitefly hosts on Gerbera jamesonii

One of the factors that may complicate biological control of the greenhouse whitefly on Gerbera jamesonii by Encarsia formosa is the rosette shape of this ornamental, which differs from the vertical shape of most vegetable plants (cucumber, egg plant, tomato, etc.). Therefore, host-habitat location and the behaviour prior to landing on uninfested and infested leaves was studied. Attraction of E. formosa from a short distance by infested leaves could not be detected: the parasitoid females landed at random on uninfested and infested leaves. After the first landing, a redistribution of the wasps occurred on the leaves. After 24 h three times as many wasps were found on the infested leaves than on uninfested ones. In a dispersal experiment with four plants, E. formosa appeared to have no preference for landing on leaves of the medium age class, which is the age class on which most of the whiteflies in a suitable stage for parasitism occur. Twenty percent of the parasitoids were found on the plants 20 min after releasing them. These results were independent of the plant cultivar and the host density on the plants. In the course of 8 h, the number of E. formosa females recovered from plants increased linearly, and this increase was greater on plants where hosts were present and also greater on the plant cultivar with the lowest trichome density. After 24 h, the percentage of females was highest (56%) on plants with the highest host density. E. formosa females were arrested on leaves where hosts were present. Contrary to our expectation, the results from the two G. jamesonii cultivars that differed strongly in leaf hairiness were not significantly different in most experiments. Only at the high host density was parasitism found to be lower on the cultivar with the higher hair density. Parasitoids may walk on top of the `hair coverlet' of cultivars with high trichome density and, therefore, be hampered less than expected.     

The role of pre- and post-imaginal experience in the host-finding and oviposition behaviour of the cabbage moth

The influence of pre- and post-imaginal experience on the orientation, landing, and oviposition of the female cabbage moth, Mamestra brassicae (L.) (Lepidoptera: Noctuidae) was studied in a wind tunnel. The results of experiments on the influence of pre-imaginal experience do not support the Hopkins host selection hypothesis because the females’ responses to host plants were not affected by the plant species used for feeding the larvae. The females were initially attracted to and landed on chrysanthemum whether or not they were fed on this plant species. In addition, the oviposition preference for cabbage plants was not changed by the larval feeding regimen. Imaginal experience influenced the females’ subsequent orientation/landing responses: exposure of the females to chrysanthemum plants for 5 min decreased their subsequent responses to this species, but subsequent responses to cabbage were not changed when the females were exposed to cabbage plants. Overnight exposure of females to a chrysanthemum or cabbage plant decreased the subsequent orientation/landing on the same species offered in non-choice tests. A similar effect was observed in responses to chrysanthemum plants after overnight contact, and possibly to cabbage plants, in the two-choice tests. Overnight exposure to cabbage or chrysanthemum had no effect on subsequent oviposition: females from both treatments oviposited more often on cabbage.     

Learning of host-infested plant volatiles in the larval parasitoid Cotesia kariyai

The ability to learn plant volatiles in Cotesia kariyai females was examined by wind tunnel bioassays. Searching experience on a host-infested corn plant increased subsequent flight responses of females to the infested plant. Females experiencing host by-products together with the volatiles extracted from infested leaves one time showed an increased response. However, such behavioral changes were not observed in females which experienced only the host by-products or the volatiles. Thus, the increased response is considered to be preference learning. Multiple experiences of C. kariyai with host by-products together with the volatiles did not increase their flight response to the volatiles. Furthermore, this learned response gradually decreased within 2 days. These behavioral modifications based on experience would be advantageous for C. kariyai to locate their polyphagous hosts efficiently.     

嗜食植物与非嗜食植物对西花蓟马成虫寄主选择性的影响

【目的】西花蓟马是世界性害虫,利用西花蓟马对寄主植物嗜食性的差异,通过驱避作用防控西花蓟马,能够为绿色治理提供依据。【方法】在西花蓟马嗜食的甘蓝和非嗜食的大蒜上互喷汁液,采用黄瓜+甘蓝、黄瓜+大蒜2种相间种植方式,研究它们对西花蓟马寄主选择性的影响。【结果】在甘蓝上喷洒大蒜汁液后,甘蓝叶片上西花蓟马的虫量和产卵量均明显减少。大蒜汁液浓度越高,减少得越多;在大蒜上喷洒甘蓝汁液后,大蒜植株上西花蓟马的虫量和产卵量明显增加,且增加程度与甘蓝汁液的浓度呈正相关。黄瓜和甘蓝相间排列时,西花蓟马在黄瓜叶片上的数量与单作黄瓜叶片上无明显差异;但黄瓜和大蒜相间排列时,西花蓟马在黄瓜上的数量明显高于单作黄瓜上的虫数,多52.4%。【结论】在嗜食寄主植物上喷洒非嗜食植物汁液或间作非嗜食的寄主可以明显减少西花蓟马的选择性。研究结果为利用非嗜食植物挥发物防控西花蓟马提供了理论依据和新的方法。     

Host odours enhance the responses of adult Rhyzopertha dominica to male-produced aggregation pheromone

The behavioural responses of adult male and female Rhyzopertha dominica (Coleoptera, Bostrichidae) to blends of host volatiles and male-produced aggregation pheromone were observed in a four-arm airflow olfactometer. The odour sources used were five pheromone-releasing males each on a single maize grain (lower maize-volatiles blend), five pheromone-releasing males on 500 g of maize (higher maize-volatiles blend) or the host volatiles emanating from 500g of maize (maize volatiles alone). Multiple-choice tests, in which volatiles from all three odour sources were presented in the exposure chamber at the same time, were used to study odour preferences of the males and females. Both sexes showed strongest attraction to the higher maize volatile blend but there were significant sex differences in response to the odour sources. Males spent significantly more time than females in the zone with only maize volatiles, and females spent significantly more time in the higher maize-volatiles zone. However, when odour sources were offered singly, females gave numerically greater responses than males to all sources although this difference was statistically significant for only the lower maize-volatiles blend. As males are more attracted than females to host odours alone it is suggested that they may be more highly adapted to seek out new hosts while females are more inclined to locate a food source by following the pheromone signals produced by males. However, both sexes responded most strongly to the odour source comprising aggregation pheromone with the higher proportion of maize volatiles.     

Host recognition by the specialist endoparasitoidMicroplitis croceipes (Hymenoptera: Braconidae): Role of host- and plant-related volatiles

The specialist parasitoidMicroplitis croceipes Cresson can parasitize only noctuid larvae in the generaHelicoverpa andHeliothis. To be successful in their search for hosts, the ability to distinguish hosts from nonhosts feeding on the same plant is beneficial. In flight tunnel experiments, we found that prior to landing on the odor sourceM. croceipes were able to distinguish volatiles released from frass of host larvae(Helicoverpa zea Boddie) and nonhost larvae (Spodoptera exigua Hübner andSpodoptera frugiperda J. E. Smith) fed on cotton. However, an initial contact experience with frass of cotton-fed host larvae appeared to be critical for this ability. Wasps that had antennated frass of host larvae fed pinto bean diet were equally attracted to frass of host and nonhost larvae fed on pinto bean diet. In short-range walking experiments, wasps located cotton-fed host larvae faster than diet-fed larvae, regardless of their experience. Wasps that had antennated frass of cotton-fed host larvae were less attracted to cotton-fed nonhost larvae, compared to host larvae, and preferred to sting host larvae. Plant-related volatiles in host frass and larvae appear to play a major role in the successful location of host larvae.     

The Role of Chemical Cues in Host and Mate Location in the Pear Psylla Cacopsylla bidens (Homoptera: Psyllidae)

Chemical communication was shown to play a role in the pear psylla, Cacopsylla bidens. Electrophysiological (EAG) and behavioral responses were investigated in males and females pear psylla . Males were found to be attracted to females, and especially to those on host plants, but not to males, uninfested host plants, or plants infested with conspecific larvae. On the other hand, females were not attracted to males or females but displayed some attraction to host plants. Furthermore, females showed a preference for uninfested pear versus plants infested with conspecific larvae. The antennae of males gave highest electroantenographic response to volatiles from pears infested with females but not males, while females, responded also toward the volatiles of pear alone. These results indicate that females of C. bidens emit sex pheromones that are attractive to the males and suggest that, host volatiles may play a role in host selection by pear psylla females.     

3种寄主上桃蚜的选择性及形态分化

桃蚜是一种重要的农业害虫,寄主广泛,种下分化复杂。以采自黄土高原旱作区桃树、烟草、甘蓝上的桃蚜为研究对象,通过叶片选择法、传统比较形态测定法研究了3种寄主上桃蚜的选择性及形态分化。结果表明:在3种寄主同时存在的情况下,烟草上的桃蚜嗜食烟草,表现为63.5%的桃蚜选择烟草叶,13.8%选择甘蓝叶,8.2%选择桃叶,而甘蓝和桃树上的桃蚜对原寄主没有表现出明显的嗜好性;从形态指标来看,3种寄主上的桃蚜在体长、触角末节长度、后足腿节长度、触角与体长的比例方面存在显著差异(P0.05),说明这几个特征可以作为区分这3种寄主上桃蚜的依据。综合分析可以初步认为黄土高原旱作区烟草上的桃蚜可能形成了寄主专化型-烟草型。     

Olfactory response of three parasitoid species (Hymenoptera: Braconidae) to volatiles of guavas infested or not with fruit fly larvae (Diptera: Tephritidae)

The olfactory responses of the native parasitoids Doryctobracon areolatus (Szépligeti) and Asobara anastrephae (Muesebeck) and of the exotic parasitoid Diachasmimorpha longicaudata (Ashmead) to guava (Psidium guajava L.) infested or not with fruit fly larvae were evaluated. D. areolatus and D. longicaudata females responded to the odors of uninfested rotting guavas, although D. areolatus was also attracted to fruits at the initial maturation (turning) stage. The females of these species recognized the volatiles of guavas containing Ceratitis capitata (Wied.) larvae. However, in bioassays involving fruits with larvae of different instars, D. longicaudata females were not able to separate between fruits containing C. capitata larvae at the initial instars and larvae at the third instar. In the evaluations of volatiles released by guavas containing C. capitata and Anastrepha fraterculus (Wied.) larvae, the D. longicaudata females were oriented toward the volatiles of fruits containing both host species, but differed significantly from volatiles of guavas containing C. capitata larvae. The D. areolatus females also showed responses to both species, although with a preference for volatiles of fruits containing A. fraterculus larvae. The A. anastrephae females were oriented toward the odors of fruits infested with both fruit fly species. In the shade house, D. longicaudata females were oriented to volatiles of rotting fruits containing larvae or not, but could not significantly differentiate between hosts. D. areolatus females were not attracted toward fruits on the ground in the shade house, regardless of host, suggesting that this parasitoid does not forage on fallen fruits.     

Oviposition behavior of the cabbage root fly,Delia radicum (L.), influenced by host plant extracts

An ethanolic extract of cabbage leaves (Brassica oleraceavar. capitata,Golden Acre)and derived fractions were tested on natural and surrogate leaves in order to study the oviposition behavior of the cabbage root fly Delia radicum(Diptera: Anthomyiidae). On surrogate plastic leaves coated with a thin layer of paraffin wax and treated with 0.1 gram leaf equivalent (gle) of an ethanolic raw cabbage extract, the females displayed the same sequence of behavioral patterns as on a natural host plant. The quantified oviposition behavior correlated well with the actual number of eggs laid. The extract-treated surrogate leaves received at least as many eggs as natural leaves with a similar surface area. Previous exposure to surrogate or natural leaves seemed not to induce a specific preference. The three fractions (hexane, butanol, and water) of the raw extract stimulated the oviposition behavior. This was taken as evidence for the presence of a multicomponent mixture or a chemical pattern in the cabbage leaves stimulating oviposition. At the tested concentration each fraction alone could stimulate in some individuals the complete behavior. No effect of volatile components of the raw extract could be detected. This is thought to be due to the extraction procedure, which limits the production of attractive, volatile compounds, such as isothiocyanates.     

Deterrent effects of intact plants on host-searching behavior of parasitoid wasps

Female parasitoids distinguish between host-infested and intact plants using chemical cues; however, the contribution of intact plants to host searching of parasitoids has not been investigated so far. Here, we tested how host-searching behavior of the parasitoid wasp, Cotesia kariyai (Watanabe) (Hymenoptera: Braconidae), was affected by intact maize plants in a wind tunnel. To determine the best color for material to create a plant model, we observed flight responses of female wasps to paper plant models of four different colors. Wasps tended to land more frequently on green models than other models. Therefore, a green paper model was used for subsequent experiments. In a no-choice test, female wasps showed higher landing rates on a paper plant model treated with herbivore-induced plant volatiles (HIPVs) than on an intact plant. Moreover, in two-choice tests, wasps preferred the plant model with HIPVs over an intact plant with HIPVs. Intact plants seem to deter C. kariyai females. Our findings suggest that information from intact plants also contributes to the host-searching behavior of females in the natural environment.

     

Feeding damage by larvae of the mustard leaf beetle deters conspecific females from oviposition and feeding

Herbivorous insects may be informed about the presence of competitors on the same host plant by a variety of cues. These cues can derive from either the competitor itself or the damaged plant. In the mustard leaf beetle Phaedon cochleariae (Coleoptera, Chrysomelidae), adults are known to be deterred from feeding and oviposition by the exocrine glandular secretion of conspecific co-occurring larvae. We hypothesised that the exocrine larval secretion released by feeding larvae may adsorb to the surface of Chinese cabbage leaves, and thus, convey the information about their former or actual presence. Further experiments tested the influence of leaves damaged by conspecific larvae, mechanically damaged leaves, larval frass and regurgitant on the oviposition and feeding behaviour of P. cochleariae. Finally, the effect of previous conspecific herbivory on larval development and larval host selection was assessed. Our results show that (epi)chrysomelidial, the major component of the exocrine secretion from P. cochleariae larvae, was detectable by GC-MS in surface extracts from leaves upon which larvae had fed. However, leaves exposed to volatiles of the larval secretion were not avoided by female P. cochleariae for feeding or oviposition. Thus, we conclude that secretion volatiles did not adsorb in sufficient amounts on the leaf surface to display deterrent activity towards adults. By contrast, gravid females avoided to feed and lay their eggs on leaves damaged by second-instar larvae for three days when compared to undamaged leaves. Mechanical damage of leaves and treatment of artificially damaged leaves with larval frass or regurgitant did not affect oviposition and feeding of P. cochleariae. Since no adverse effects of previous herbivory on larval development were detected, we suggest that female P. cochleariae avoid Chinese cabbage leaves damaged by feeding larvae for other reasons than escape from competition or avoidance of direct negative effects that result from consuming induced plant material.     

Role of visual cues and interaction with host odour during the host-finding behaviour of the cabbage moth

The approach and landing responses of female Mamestra brassicae (L.) (Lepidoptera: Noctuidae) to visual cues from artificial plant leaves of different shapes and presence/absence of cabbage plant odour were investigated in a laboratory wind tunnel. The leaves were painted with cadmium yellow colour and observed under dim red light. Females showed oriented flight towards plant odours but landed significantly more often when the odour was presented with an artificial leaf. In three-choice tests, the shape of the leaf targets (circle, square or triangle) did not influence the female response. However, the size of the target did influence the insect response: the females preferred landing on square targets with sides of 5 or 10 cm rather than on the largest target, with sides of 15 cm. The orientation of the target influenced the insects' response: females landed significantly more often on the target positioned vertically than horizontally.     

Generalism Versus Specialism: Responses of Diadegma mollipla (Holmgren) and Diadegma semiclausum (Hellen), to the Host Shift of the Diamondback Moth (Plutella xylostella L.) to Peas

A population of the diamondback moth Plutella xylostella (L.) (Lepidoptera: Plutellidae) (DBM) was recently found to infest sugar snap- and snowpeas in the Rift Valley in Kenya, causing heavy damage. The influence of this host shift on host location preferences of two parasitoids was investigated: The indigenous Diadegma mollipla (Holmgren) regarded as a relative generalist, and Diadegma semiclausum(Hellen), regarded as highly specific to DBM. The attractiveness of different odour sources was compared for the two parasitoid species using a Y-tube olfactometer using naïve females. D. mollipla was not significantly attracted to any cabbage related odours but showed a significant preference for the DBM infested pea plant when tested against clean air. D. semiclausum was highly attracted to the undamaged cabbage plant and odours related to cabbage. On the other hand, peas infested with DBM, showed no attractiveness to this parasitoid. The results showed that specialisation of D. semiclausum is mediated by host plant signals, associated with crucifers, which are not encountered in DBM feeding on peas. For D. mollipla,although a frequent parasitoid on DBM in crucifers, volatiles emitted by these plants might not be used as primary cues for host location. This species may respond largely to chemicals yet unknown and associated with a variety of plant-herbivore interactions.     

Influence of visual stimuli on host location in the butterfly, Eurema hecabe

The influence of visual stimuli on female host location was investigated in Eurema hecabe (Lepidoptera: Pieridae). Mated females with no oviposition experience were presented with artificial plant models treated with the methanol extract of the host plant, Lespedeza cuneata (Fabaceae). When models of different colours were presented to females, they landed and deposited eggs predominantly on the yellow-green model. However, the size of the model was not important. When females were presented with simple square models and a model on which small squares were assembled, they predominantly chose the assembly model. This suggests that female discern the pattern which resembles the leaf of their main host plants, which consists of many small leaflets. However, the whole image of a leaf of the host plant was not an essential cue for host location, as females also chose the square with the central space more frequently than that without the space. We suggest that the long contour of a complicated form plays a role in female host location.     

Effects of recent experience on foraging in tephritid fruit flies

In field-cage studies, we investigated how the foraging behavior of tephritid fruit flies is modified by experience immediately prior to release on host plants. We observed females of a relatively monophagous species,Rhagoletis mendax (blueberry maggot fly), an oligophagous species,Rhagoletis pomomella (apple maggot fly), and a polyphagous species,Ceratitis capitata (Mediterranean fruit fly). Just prior to release on a host plant, the following kinds of stimuli were supplied: (1) single oviposition in a host fruit, (2) contact with 20% sucrose, (3) contact with a mixture of protein food (bird feces and sucrose), (4) contact with water, and (5) a walk over a host-plant leaf. When flies foraged on host plants without resources, search was most intensive (as measured by number of leaves visited) following a single oviposition in fruit, but residence time generally was the same following exposure to sugar, protein, and fruit stimuli.Rhagoletis mendax andC. capitata females visited the fewest leaves following exposure to water or host leaves, whereasR. pomonella foraged equally intensively following exposure to food stimuli, water, or leaves. On host plants containing resources (fruit and protein food), a single oviposition dramatically increased the number of females of all three species that found fruit compared to females that received experience with food, water or foliar stimuli. We found no significant effect of recent brief experience with any of the stimuli on subsequent attraction to protein food. Overall,C. capitata exhibited a higher propensity to abandon host plants than eitherR. mendax orR. pomonella. We suggest that this may reflect adaptations to differences in distribution of host plants in nature, strategies of dispersal, and host range.