Electroantennogram and behavioural responses of different forms of the bird cherry-oat aphid,Rhopalosiphum padi,to sex pheromone and a plant volatile

Winged (alate) virginoparae were induced in the laboratory by crowding the bird cherry-oat aphid, Rhopalosiphum padi (L.) under long-day conditions. Males and gynoparae (the winged female form that produces sexual females) were induced by short days. Electroantennogram (EAG) and behavioural responses were investigated in each of the three forms to two aphid sex pheromone components, (-)-(4aS,7S,7aR)-nepetalactol and (+)-(4aS,7S,7aR)-nepetalactone, and benzaldehyde, a volatile which is released by the winter host plant. All three compounds elicited EAG responses with the males showing the highest sensitivity to each compound. Both the nepetalactol and the nepetalactone elicited larger EAG responses in gynoparae than in the winged virginoparae but antennae from virginoparae were more responsive to benzaldehyde. Although the nepetalactone is not a sex pheromone component in R. padi the EAG responses were similar to those evoked by the nepetalactol, the sex pheromone, in all three aphid forms. In a linear-track olfactometer, significantly more male R. padi moved into air containing nepetalactol, nepetalactone or benzaldehyde than into a simultaneous choice of clean air (i.e. attraction) but nepetalactol was more attractive than nepetalactone. Males, however, showed no response to a mixture of nepetalactol and nepetalactone. Gynoparae were attracted only to the nepetalactol but were less sensitive than the males and showed no response to the nepetalactone or benzaldehyde. In contrast, alate virginoparae showed no behavioural responses to any of the compounds. The present study supports the idea that the male R. padi utilise both sex pheromone and benzaldehyde for mate/host-plant location in autumn. It also demonstrates, for the first time, polyphenic differences in the olfactory responses at the peripheral level between the two female forms. Such differences impact on the life-cycle strategy where winged virginoparae move between graminaceous summer host plants while gynoparae move from the summer hosts to the bird cherry, winter host. The latter move appears to be assisted by the sex pheromone released by sexual females, already present on that host, acting as an aggregation pheromone.     

蚜虫雄蚜与雌性母对性信息素及植物挥发物的田间反应

田间观察了桃蚜Myzus persicae (Sulzer)、绣线菊蚜Aphis spiraecola Patch 、山楂圆疣蚜Ovatus crataegarius (Walker)等3种蚜虫对性信息素[(4aS,7S,7aR)-荆芥内酯和 (1R,4aS,7S,7aR)-荆芥醇]的反应,并且调查了性信息素与植物挥发物对桃蚜的田间引诱活性的相互作用.在有冬寄主或夏寄主植物的田中,性信息素诱捕器诱捕到桃蚜雄蚜与雌性母的数量显著多于对照诱捕器的诱捕数;但在非寄主植物的田中,却引诱不到桃蚜.苯甲醛(冬寄主植物桃树Prunus persica的主要挥发物组分之一)能够增强桃蚜雄蚜的引诱作用.绣线菊蚜雄蚜和雌性母对植物中提取的荆芥内酯有反应,而山楂圆疣蚜雄蚜和雌性母对植物中提取的和人工合成的荆芥内酯都没有反应,但对荆芥醇有反应.并且在荆芥醇中添加荆芥内酯之后对山楂圆疣蚜雄蚜引诱活性显著提高.还讨论了雌性蚜产生化合物被雄蚜作为性信息素、被雌性母作为聚集信息素以及性信息素与寄主植物挥发物的相互作用.     

Perception of volatiles associated with sex and food by different adult forms of the black bean aphid, Aphis fabae

Abstract. Electroantennograms (EAGs) were recorded from adult male and asexual forms (winged and wingless virginoparae and gynoparae) of the black bean aphid, Aphis fabae , during stimulation with two sex pheromone components, (+)-(4a S ,7 S ,7a R )-nepetalactone and (-)-(1 R ,4a S ,7 S ,7a R )-nepetalactol, as well as six plant volatiles, i.e. ( E )-2-hexenal, ( E )-2-hexenol-1, ( Z )-3-hexenol-1, ( Z )-3-hexenyl acetate, hexanal and allyl isothiocyanate. The male antennae are 1000-10,000 times more sensitive to nepetalactol and nepetalactone than to the plant compound ( E )-2-hexenal. Besides this marked difference of EAG peak responses in males, the EAG rise and decay are slower for the sex pheromone components. Males are also much more sensitive to the sex pheromone components than asexual females. This high sensitivity correlates with a predominance of antennal secondary rhinaria, the major sites of pheromone perception in the male. However, it is the primary rhinaria on the antennae of the wingless asexual females that are responsible for pheromone perception. Male antennae are as responsive as the asexual female antennae to the plant volatiles. The specialization of the male for mate location is discussed.     

Functional specialisation and polyphenism in aphid olfactory sensilla

Electroantennogram (EAG) responses to the aphid sex pheromone components, (1R,4aS,7S,7aR)-nepetalactol and (4aS,7S,7aR)-nepetalactone, and a plant volatile, (E)-2-hexenal, were investigated at three different positions (5/6th, 4/5th and 3/4th inter-segmental regions) along the antennae of four different morphs in two host-alternating aphid species, Aphis fabae Scopoli and Rhopalosiphum padi (L.). Position-dependent and morph-specific EAG responses were elicited in both species. The nepetalactol and nepetalactone isomers elicited large EAG responses at all three recording positions in males of both species, such that primary rhinaria as well as secondary rhinaria appeared to respond. Asexual female morphs showed relatively smaller EAG responses to these compounds. The secondary rhinaria, which have been reported as sex pheromone receptors in males, were not very different in their number and distribution between gynoparae and alate virginoparae, but the gynoparae showed significantly larger EAG responses to nepetalactol and nepetalactone. The alate virginoparae showed EAG responses that were similar to those of apterous virginoparae, which lack secondary rhinaria. Taking the EAG response profiles together with the distribution of the secondary rhinaria, it is suggested that the function of secondary rhinaria differs between the morphs. Secondary rhinaria appear to detect sex pheromone components in males and gynoparae but not in the alate virginoparae. If they are functional in the latter morph, they are likely to play a role in detecting specific, but as yet unknown, volatile compounds. Some 30 plant volatiles were tested but none evoked an EAG response that could be allocated to the secondary rhinaria. In contrast to the very different EAG response profiles to the pheromone compounds between morphs, EAG responses to (E)-2-hexenal were similar in all forms and both species. These findings suggest that this plant volatile was detected only by the two primary rhinaria, which are common to all morphs. The present study showed that EAG responses were not a simple summation of receptor potentials between recording and reference electrodes in aphids. The localised distribution pattern of olfactory receptor neurones around the recording electrode was also likely to contribute to the EAG responses.     

蚜虫性信息素对桃园蚜虫的引诱作用

将从荆芥Nepeta cataria L.中提取的纯度为96%的荆芥内酯和通过二异丁基铝还原提取的荆芥内酯得到的纯度为67%荆芥醇,配成浓度为5%的不同比例的己烷溶液,于1995年10月19～28日在北京大兴县南郊农场的桃园进行了引诱蚜虫的试验。各种比例的性信息素诱捕器均能引诱到桃蚜Myzus persicae (Sulzer)雄蚜、桃粉大尾蚜Hyaloptera amygdali Blanchard雄蚜以及少量的桃蚜产雌性母;荆芥醇和荆芥内酯的比例为1∶1时引诱桃蚜雄蚜的效果最好;单一组分的荆芥醇引诱桃粉大尾蚜雄蚜的效果最好。     

豌豆蚜触角转录组化学感受蛋白的鉴定、表达和结合特性分析

[目的]本研究旨在鉴定豌豆蚜Acyrthosiphon pisum触角转录组中化学感受蛋白(chemosensory protein,CSP)基因,明确触角中高表达的豌豆蚜CSP蛋白与蚜虫报警信息素、性信息素以及植物挥发物的分子结合特性.[方法]通过对豌豆蚜成蚜触角进行转录组测序,鉴定触角中候选CSP基因;采用RPKM...     

Electrophysiological characterisation of olfactory sensilla in the black bean aphid, Aphis fabae

Electrophysiological responses of three different olfactory sensilla (proximal primary rhinaria (PPR), distal primary rhinaria (DPR) and secondary rhinaria (SR)) to the sex pheromone components, nepetalactol, nepetalactone, and a plant volatile, (E)-2-hexenal, were investigated in four different morphs of the black bean aphid, Aphis fabae Scopoli. The DC responses recorded directly from the antennal sensilla and termed electrosensillograms (ESGs) were generally much larger (up to 14 mV) than electroantennogram (EAG) responses (up to 2.5 mV). Characteristic morph-specific response profiles to these compounds were observed in each type of rhinarium and response waveforms were different between (E)-2-hexenal and the sex pheromone components. (E)-2-Hexenal elicited the largest responses at PPR, while nepetalactol and nepetalactone elicited the largest responses at SR in gynoparae and males. Nepetalactol and nepetalactone also showed significant activities on DPR and PPR in all morphs. In contrast, (E)-2-hexenal had almost no activity on SR. However, almost all of the SR investigated in males and gynoparae were sensitive to both nepetalactol and nepetalactone. A small sub-set of male SR responded mainly to nepetalactol. SR of winged virginoparae did not respond to the sex pheromone compounds. Paraffin oil (the solvent control) also elicited significant responses at PPR in virginoparae but not in other morphs. In a further experiment, SR of winged virginoparae showed no response to 30 other plant volatile compounds or the alarm pheromone component, (E)-beta-farnesene. Nepetalactol and nepetalactone had similar dose-response profiles in the SR of gynoparae. The results indicate that SR in males and gynoparae are highly specialised to detect sex pheromone compounds, while the DPR and PPR are relatively broadly tuned to both plant volatiles and sex pheromone components. The presence of SR in winged virginoparae that are not responsive to sex pheromone components, alarm pheromone, or any of the plant volatile compounds tested may indicate a possible role of these sensilla to detect, as yet, unknown compound(s) with a high specificity. The present study also suggests that PPR may play a role in detecting paraffin-related compounds such as cuticular hydrocarbons of plants.     

Aphid sex pheromones: from discovery to commercial production

This review charts the progress made with aphid sex pheromone chemistry, from initial identification of cyclopentanoid nepetalactones, for example (4aS,7S,7aR)-nepetalactone (1). and (1R,4aS,7S,7aR)-nepetalactol (2). to commercial production from a renewable non-food crop, the catmint, Nepeta cataria (Lamiaceae). The availability of aphid sex pheromone components is now facilitating the development of new aphid pest control strategies, incorporating the use of other semiochemicals, particularly in the manipulation of populations of aphid parasitoids and aphid predators such as lacewings, which can utilise the nepetalactones and closely related molecules to locate their hosts and prey. This is the first example of a plant resource being developed as a feedstock for the production of a commercially valuable insect pheromone. The development of a plant-based production route highlights the tremendous potential that higher plants offer as cheap and renewable resources for the production of insect semiochemicals, through the wide array of secondary metabolites that they can generate.     

Cereal aphid responses to sex pheromones and host-plant odours in the laboratory

Abstract. The movement of Sitobion fragariae (Wlk.) (Homoptera: Aphididae) males towards odours from conspecific sexual females (oviparae) and the aphid sex pheromone component (4a S ,7 S ,7a R )-nepetalactone were demonstrated in a linear-track olfactometer. Males of the closely related Sitobion avenae (Fab.) were attracted to odours from conspecific oviparae and also oviparae of S. fragariae. These results were consistent with the recent identification of this nepetalactone isomer as the major component of the sex pheromone in these species. Males of S.fragariae were not significantly attracted to oviparae of S.avenae. indicating some qualitative or quantitative difference in the pheromones of the two species. Males did not respond to the related (1R,4aS,7S,7aR)-nepetalactol or to host-plant odours. Gynoparae of these species moved towards the nepetalactone and also to host-plant odours. A combination of both stimuli was more attractive than plant odour alone. Gynoparae of S.fragariae also responded significantly to the combined treatment when this was tested against the nepetalactone.     

蚜虫性信息素的田间引诱蚜虫试验

据报道蚜虫的性信息素的主要成分是（＋）－（４ａＳ,７Ｓ,７ａＲ）－荆芥内醌（１）和（＋）－（１Ｒ,４ａＳ,７Ｓ,７ａＲ）＝荆芥醇（Ⅱ）。应用这两种物质进行引诱雄性蚜的田间实验,取得了良好的效果,初步获得了应用于田间的合适的配比及使用的时间。     

Sex pheromone and visual trap interactions in mate location strategies and aggregation by host-alternating aphids in the field

Abstract. Field observations were made on the responses of males and gynoparae of three host-alternating aphid species, the blackberry-cereal aphid, Sitobion fragariae (Walker), the bird cherry-oat aphid, Rhopalosiphum padi (L.) and the damson-hop aphid, Phorodon humuli (Schrank) to species-specific sex pheromones released from transparent and coloured water traps.Pheromone traps caught significantly more males than did control traps without pheromone, whereas transparent, light green, yellow and orange traps caught most insects.Measurements of the distance over which sex pheromones function indicated that male P.humuli detect the pheromone 2–6 m from the source and can fly upwind to a source in wind speeds of 0.7 m s^-1.In all three species significantly more gynoparae were caught in pheromone traps than in control traps, suggesting that pheromone released by adult sexual females may assist late-flying gynoparae to locate a suitable host plant on which to deposit their progeny.The response is relatively stronger for males than gynoparae, but the pheromones appear to act as both sex and aggregation pheromones.     

蚜虫性信息素

蚜虫属同翅目蚜总科,是重要的农作物害虫。蚜虫长年行孤雌生殖,在中、高纬度深秋时,产生性蚜,以卵越冬。雌性蚜分泌性信息素荆芥内酯(nepetalactone)和荆芥醇(nepetalactol)。该文综述了蚜虫性信息素的结构与组分、性腺的形态结构、性信息素释放及影响因素;性信息素对雌性母和天敌的引诱作用;性信息素在蚜虫种间隔离中的地位和作用以及蚜虫性信息素在蚜虫防治中的应用前景。     

Flight behaviour of the bird cherry aphid, Rhopalosiphum padi

Abstract Seasonal winged forms of the bird cherry aphid, Rhopalosiphum padi L., were flown in an automated flight chamber. During their maiden flights, autumn migrants (gynoparae) had a greater initial rate of climb than summer (exules) and spring migrants (emigrants). Autumn migrants, presented with a laterally-positioned green-illuminated target for 2.5 s during each minute of flight, flew for a longer period before responding to the target (110 min) than summer migrants (36 min), while spring migrants responded after an intermediate time (72 min). This period is equivalent to the migratory phase of flight. The summer migrants were maximally responsive to targets illuminated by green monochromatic light (X, 555 nm). Targets illuminated with ultraviolet (γλ 360 nm) were attractive during the migratory and green-responsive phases of flight, but infrared (γλ700 nm) wavelengths were not attractive. The results are discussed with respect to aphid behaviour in the field.     

Pheromone Traps for the Dissemination of an Entomopathogen by the Damson-hop Aphid Phorodon humuli

Two types of sex pheromone-emitting trap, a simple louvred trap and a commercially-produced wasp trap (Waspy®) designed or modified as delivery systems for entomopathogens, were tested using an isolate of the pathogen Verticillium lecanii . Their efficiencies for capturing winged autumn females (gynoparae) and males of Phorodon humuli in a hop garden and a plum orchard were compared with each other and with catches by yellow-painted water traps. Gynoparae rarely entered either type of pathogen dissemination trap. Males only entered traps that released (1 R , S ,4a R , 7 S , 7a S )-nepetalactol, the sex pheromone of P. humuli . In the hop garden, approximately five times as many males entered the modified Waspy® as entered the louvred traps, whereas in the plum orchard similar numbers of males were caught in both traps. On average, males visited the modified Waspy® trap for 34.1 ±7.9 (SE) min. Males spent about 50% of their time in the arena where the entomopathogen would normally be present and a further 20% of the time on the inside of the arena lid alongside the pheromone source. Out of a sample of 16 live aphids removed from a trap dispensing V. lecanii , 9 died from infection by the pathogen and 5 of the 16 initiated colonies of the fungus after they walked on a sterile agar plate for five minutes. None of the 15 aphids collected from traps without V. lecanii became infected or initiated colonies of the fungus on agar plates.     

The responses ofPraon spp. parasitoids to aphid sex pheromone components in the field

In autumn 1991, aphid parasitoids of the genusPraon (Hymenoptera; Braconidae) were caught in water traps with lures containing synthetic aphid sex pheromone components at three sites in England and one in Germany. At two of the English sites and at the German site, the traps were placed in winter cereal fields whilst the third English site was in woodland. Three species were caught,P. volucre, P. dorsale andP. abjectum. Those caught in cereal fields were almost entirelyP. volucre, whilstP. dorsale dominated at the woodland site. Of the known aphid sex pheromone components, the most effective lure was the (+)-(4aS, 7S, 7aR)-nepetalactone. Nepetalactone traps placed at the woodland site in spring and summer caught fewPraon females, and attraction may be confined to the autumn, when sexual female aphids are present in the field. Male parasitoids did not respond to the aphid pheromones at any time, although they were caught in suction traps operated at the woodland site during the autumn. At the cereal sites height had a significant influence on the efficiency of the pheromone traps, those placed just above the crop canopy being most effective. There was no evidence that any other genus of parasitoid responded to aphid sex pheromones at these sites.     

A transgenerational interval timer inhibits unseasonal sexual morph production in damson-hop aphid, Phorodon humuli

Abstract.  The induction of sexual and parthenogenetic morphs of the damson-hop aphid, Phorodon humuli , on hops is controlled by daylength. The ability of P. humuli , to produce winged pre-sexual females (gynoparae) in the short-day conditions of spring is inhibited by an interval timer present in generations immediately after hatching of the overwintering egg. The inhibition expires after three generations when nymphs are born and reared in short days (LD 12 : 12 h), irrespective of whether their parents are reared in short or long days (LD 18 : 6 h). No gynoparae are produced by aphids maintained for 13 generations in long days. Two wingless aphids from 35 survive transfer from Prunus spinosa to hops. No winged females are produced during nine generations among their progeny maintained in long days on hops, but gynoparae, followed by males, are produced one generation after these aphids are transferred to short days.     

Electroantennogram responses of Apanteles obliquae (Hym., Braconidae) to various infochemicals

Abstract:   The electroantennogram recording technique (EAG) was used to study the olfactory sensitivity of Apanteles obliquae (Hym., Braconidae), a gregarious larval endoparasitoid of Spilosoma obliqua (Walker) (Lep., Arctiidae), to 25 general plant volatiles belonging to alcohol, aldehyde and terpenoid groups and also to volatiles from the host and plant–host complex. The EAG data indicated different olfactory sensitivity between the sexes, not only to individual plant volatiles but also to the volatiles from host and plant–host complex. Females were found to be more responsive than males. However the synthetic sex pheromone blend of the host insect elicited similar EAG responses in both sexes. The EAG data of the present study is correlated with the reported behaviour observed in other parasitoids.     

绿盲蝽对性信息素类似物和植物挥发物的触角电位反应

为了更好地利用性信息素类似物与植物挥发物对绿盲蝽Lygus lucorum进行监测和生物防治, 本研究采用电生理学方法测试了绿盲蝽成虫对9种性信息素类似物和12种植物挥发性物质的触角电位(electroantennogram, EAG)反应。结果表明: 在测试的这21种化合物中, 绿盲蝽对丁酸酯类化合物和绿叶气味物质的EAG反应较强; 雄虫对性信息素类似物比雌虫更敏感, 而雌虫对植物挥发物更敏感。其中性信息素类似物中, 绿盲蝽对反-2-丁酸己烯酯的EAG反应最强; 植物挥发物中, 绿盲蝽对反-2-己烯醛的EAG反应最强, 雌虫对反-2-己烯醇、反-2-己烯醛、顺-3-己烯醇等绿叶气味物质的EAG反应相对高于萜类化合物的反应。剂量反应曲线表明, 在测定的浓度范围内, 雄虫对反-2-丁酸己烯酯的反应比雌虫显著高, 而对β 蒎烯的反应则比雌虫显著低。绿盲蝽成虫对性信息素类似物和植物挥发物的EAG反应存在的两性差异具有生态适应意义: 雄虫对性信息素类似物更敏感将更有利于其寻找交配对象, 而雌虫对绿叶气味物质更敏感将更有利于其寻找产卵寄主。     

Migratory and targeted flight in seasonal forms of the black bean aphid, Aphis fabae

ABSTRACT. Adult winged forms of Aphis fabae Scop., the spring migrants (fundatrigeniae; alate virginoparae), the summer migrants (alate virginoparae) and the autumn migrants (gynoparae), were flown in an automated vertical wind tunnel, with a small green target presented every 60 s. On average, targeted flight developed in the spring and summer migrants after 19 and 15 min, respectively. However, in autumn migrants targeted flight occurred after 184 min of flight. No evidence of target approach was found up to 30 min prior to this time, although some movement towards the target was observed in the 30 min before the first response of autumn migrants, suggesting that a gradual change from migratory to targeted flight occurred. No correlation was found between aphid weight and the time to first response for spring or summer migrants, but a positive correlation was found for autumn migrants. Flight patterns of autumn migrants, in the absence of a target, are described and compared with flight patterns previously reported for summer migrants.     

豆野螟成虫触角对植物挥发性和性信息素化合物的触角电位反应

以顺3-己烯乙酸酯为标准化合物,研究了豆野螟雌、雄成虫触角对不同浓度的植物挥发性和性信息素化合物组分的触角电位（EAG）反应.结果表明：豆野螟雌、雄成虫触角对植物挥发性气味具有强烈的EAG反应,但在大部分浓度下,雌、雄虫的反应没有显著差异.在测试的9种植物挥发性化合物中,雌、雄虫对高浓度反-2-己烯醛刺激的相对EAG反应的最大值分别达到250%和260%,且雌、雄虫触角对这种化合物的EAG反应的形状也有很大差异;但对豇豆花气味中的柏木烯和柏木脑没有明显的EAG反应.雌、雄蛾对性信息素都有明显反应,且雌、雄虫的反应存在显著差异,雄虫触角对性信息素化合物主要组分（反10,反12）-十六碳二烯醛的EAG反应的最大值达到250%,远超过雌虫反应的最大值(53%).雌、雄成虫这种神经感觉反应的差异反映了它们在感器结构、功能和行为上的差异.