The influence of parasitism on wing development in male and female pea aphids

Wing formation in presumptive alate morphs (virginoparae and males) was observed for the pea aphid, Acyrthosiphon pisum, exposed to attack by the parasitoid, Aphidius ervi, at different stages of host development. Morphological abnormalities in parasitized aphids such as complete apterization (development of a wingless form), formation of rudimentary wing buds, and deformed wings indicate a possible disruption of the endocrine system. Changes in the body shape and the number of olfactory secondary rhinaria on the antennae could indicate an influence of juvenile hormone in parasitized A. pisum but the development of fifth-stadium supernumerary larvae (indicated by an extra moult and which can be induced by exogenous juvenile hormone treatments) was not found in parasitized aphids. In addition, while apterization of virginoparae can also be induced by the pro-allatocidal compound Precocene III, this was not possible in the male. Males which survived parasitoid attack without forming aphid mummies (indicating that oviposition had not occurred) developed as wingless individuals suggesting that the reproductive-tract-fluids from the female parasitoid were important in the wing inhibition process. Teratocytes from the parasitoid appeared to promote developmental arrest in parasitized aphids.     

HOST ALTERNATION IN APHIS FABAE SCOP.

Experiments on the feeding preferences and relative fecundity of Aphis fabae among leaves of different ages and kinds were extended to the comparison of different forms of the aphid: apterous and alate virginoparae and gynoparae.
All three forms showed a preference for the primary host, Euonymus , over a secondary host, sugar beet, but this preference was strongest in the gynoparae, weaker in the alate virginoparae and weakest in the apterae. The relative fecundity of the aphids on the two kinds of leaf paralleled their feeding preferences most closely in the gynoparae, less so in the alate virginoparae and least in the apterae.
All three forms also showed some preference for growing over mature leaves of the same kind, but this preference was strongest in the apterous virginoparae, weaker in the alate virginoparae and weakest in the gynoparae. The relative fecundity of the aphids on the two ages of leaf paralleled their feeding preferences most closely in the apterae, less so in the alate virginoparae and least in the gynoparae.
The physiological, ecological and evolutionary significance of the results is discussed in the light of the dual discrimination theory of aphid host selection.     

Embryogenesis and oögenesis in alate virginoparae,gynoparae, and oviparae of the aphid Myzus persicae, in relation to photoperiod

The study reports on the effects of prenatal and/or postnatal exposures to short-night or long-night conditions, and of crowding, on embryogenesis and oögenesis in alate virginoparae, gynoparae, and oviparae of a holocyclic strain of the green peach aphid, Myzus persicae, from Yakima, Washington State.In alate virginoparae raised at a density of 10–20 per radish seedling in a short-night regime (8 hr darkness per diem), 3–4 embryos occurred in each of their 10 ovarioles, when the aphids attained adulthood. More than 30 larvae were deposited by most of these alatae. However, in young adult gynoparae, raised at these densities in a long-night regime (15 hr darkness per diem), only one viable embryo (a presumptive ovipara) occurred per ovariole. The follicle containing this embryo was followed by 1–2 abnormal follicles in each ovariole, and the number of larvae deposited by a gynopara was generally less than 10. In young adult oviparae similarly raised under a long-night regime, only one egg typically occurred in each of their 10 ovarioles, and the eggs deposited by an ovipara (only after it had mated) generally numbered less than 10. Alate virginoparae and gynoparae contained an additional embryo in some of their ovarioles when these morphs were raised at a lower density (1–5 per plant).Presumptive gynoparae partially developed the reproductive features of alate virginoparae when transferred to a short-night regime at birth; the converse was true when presumptive alate virginoparae were transferred to a long-night regime early in larval life. Oviparae maintained in short nights from before birth developed the appearance of apterous virginoparae but still produced eggs rather than embryos. However, their oögenesis was enhanced and eggs (10–20) were deposited by them without prior mating. Under all regimes tested, oviparae were always deposited early in the larviposition sequence of their alate mothers, and the number of oviparae deposited never exceeded 15.The possible involvement of juvenile hormone in the regulation of these events and the ecological significance of the results are 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.     

A morphological study on autumnal morphs of Aphis glycines (Hemiptera: Aphididae)

The soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), is one of the most important pests of soybean. The complex life cycle of A. glycines is characterized as heteroecious and holocyclic, and has seasonal polymorphisms occurring during its life cycle. In the autumn, A. glycines occur as winged and wingless virginoparae in soybeans, as gynoparae and males that migrate from soybeans to Rhamnus spp. and as gynoparae, males and oviparae on Rhamnus spp. In this study, wingless virginoparae, gynoparae, males and oviparae of A. glycines were successfully induced in the laboratory and morphological parameters of these morphs were selected for quantitation. To aid in identification of these A. glycines autumnal morphs, these aphids were imaged by microscope and significant differences in morphological characteristics were found: distal parts of hind leg femurs of gynoparae were grayish black, which were darker than those of winged virginoparae; the 4th instars of gynoparae, males and winged virginoparae differ greatly in dorsal abdomen coloration and covering. Our results provide an important guide for distinguishing adults of gynoparae and winged virginoparae, and for identifying nymphs of gynoparae, males and winged virginoparae.     

Production of sexual morphs by apterous virginoparae of Myzus persicae (Hemiptera: Aphididae) in relation to pre- and postnatal exposure to short day conditions

The production of males and females by apterae of a holocyclic clone of Myzus persicae (Sulzer) was studied in the laboratory by transferring apterous virginoparae from long day to short day conditions at different stages of their pre- and post-natal development. Prenatal exposure was also examined in nine additional clones coming from different regions of Greece. By increasing the number of short day cycles at 17 degrees C, the aphids switched to male production earlier and produced more males. The highest mean number of male progeny was observed in apterae that were transferred to short day conditions 13-14 days prenatally. Apterae that were transferred to short day conditions after the fourth nymphal instar produced only females. Apterous virginoparae were produced by aphids that were transferred to short day conditions 0-1 days prenatally or postnatally. Aphids, given short days prenatally or in the first instar, produced alate female progeny that were all gynoparae. The later the instar aphids were transferred to short day conditions the fewer the gynoparae produced among their female progeny. A reduction in the total number of progeny and embryos per ovariole was observed as aphids were transferred to short day conditions in earlier stages of their development. The increase in male production, which follows the increase in the duration of long night exposure, was also observed in the additional eight out of nine examined clones.     

Morph determination in the bird cherry-oat aphid, Rhopalosiphum padi L

Both crowding and poor nutrition induce the appearance of emigrants in Rhopalosiphum padi L. No emigrants developed when the aphid was reared in isolation for six successive generations on actively growing leaves of bird cherry. However, emigrants developed as soon as the leaves ceased to grow or when the aphids became crowded. Crowding of mothers and postnatal crowding of the nymphs of apterous exules both influenced the induction of alate exules. The highest proportion of alate exules developed when both mother and offspring were crowded. Short day-length induced the appearance of gynoparae and males. For 50% production of gynoparae and males, over the range of temperature 10–18d?C, a 1.75 h reduction in day-length is required for every 4d? increase in temperature. At 18 d?C, even at day-lengths as short as 6 h, exposure for three generations is required before all offspring become gynoparae or males. Low temperature and short day-length was ineffective in inducing the development of gynoparae or males in the first three generations developing from the fundatrix.     

Lipid composition in long-day and short-day forms of the black bean aphid,Aphis fabae

Long-day reared winged (alate) virginoparae from laboratory stock cultures which had been reared throughout larval development on bean stipules were significantly smaller (0.46+/-0.02 mg; mean+/-SEM) than short-day-reared gynoparae (0.69+/-0.04 mg; the winged autumn migrant) which completed development on intact beans. When winged virginoparae were raised from the third stadium on bean seedlings they grew larger (0.86+/-0.02 mg) but the gynoparae contained proportionally more total lipid (12.1+/-0.4%, gynoparae; 7.4+/-0.6%, stipule-reared virginoparae; 9.2+/-0.8%, seedling-reared virginoparae). Wingless aphids (apterae) were heavier, whether reared in short (0.99+/-0.03 mg) or long days (0.95+/-0.04 mg) but the lipid content was low (4.5+/-0.7% and 4.9+/-0.6%, respectively). The triacyl-, diacylglycerol and the phospholipid contents followed this trend but analysis of the fatty acid moieties of the triacylglycerides showed phenotypic differences. The ratios of myristic acid:palmitic acid were significantly higher in the winged forms than the wingless forms and were much higher in the gynoparae than the winged virginoparae. Short-day-reared wingless females also had a higher myristic acid:palmitic acid ratio than long-day-reared apterae, possibly reflecting the embryonic gynoparae maturing in their ovaries.     

Reproductive, morphological and behavioural affinities between the alate gynopara and virginopara of the aphid, Aphis fabae

ABSTRACT. 'Normal' alate virginoparae reared under long day conditions (LD 16:8) with postnatal crowding, and 'normal' gynoparae reared in un-crowded, short day conditions (LD 12:12) over two generations were compared in respect of the morph of the progeny, the frequency distribution of secondary rhinaria (placoid sensilla) on the third and fourth antennal segments, and preferred larviposition sites. Although in Aphis fabae the two morphs closely resemble one another morphologically, they differed in each of the above characters. Progeny sequences were analysed after transfer to the opposite photoperiodic regime at various times during postnatal development of the alate mothers. Transfer of presumptive gynoparae from short to long days induced the appearance of virginoparous progeny even when the transfer was made just after the final moult. Transfer of presumptive alate virginoparae from long to short days was effective in inducing oviparous offspring, only if the transfer was made during the first instar; oviparae were always first born. Intermediates between oviparae and viviparae were found at the time when the morph of the progeny changed. Transfer from long to short days also induced gynopara production but there appeared to be an intrinsic mechanism which prevented exclusive gynopara production by alates. When first instar, presumptive gynoparae were transferred to long days, the resulting adults would be described both reproductively and behaviourally as alate virginoparae; the opposite switch, of presumptive alate virginoparae to short days, resulted in adult gynoparae on the same criteria. However, the numbers of secondary rhinaria were influenced more by the prenatal than by the postnatal photoperiodic regime. There was an asymmetry of response in that a switch from short to long days had a greater effect on all three characters monitored than did the opposite switch from long to short days.     

Precocious metamorphosis of the aphid Myzus persicae induced by the precocene analogue 6-methoxy-7-ethoxy-2,2-dimethylchromene

For up to three days after being treated with the precocene analogue 6-methoxy-7-ethoxy-2,2-dimethylchromene, adult green peach aphids (Myzus persicae) gave birth to offspring (alate and apterous virginoparae, males) which underwent precocious metamorphosis. Only a few precocious gynoparae and no precocious oviparae were recorded. The precocious aphids became adultoid in the third or fourth-instar as a result of corpus allatum destruction. They were able to develop mature embryos, but could not deposit them because of incomplete differentiation of the reproductive tract. The treatment did not induce the production of alate virginoparae in the experimental clone of Myzus persicae. However, a few males were born late in the reproductive sequence of treated apterae. The corpora allata of the treated adults appeared on histological examination to be unaffected by precocene.     

Host-selection behaviour by genetically identical aphids with different plant preferences

The behaviour of summer and autumn winged forms of the black bean aphid, Aphis fabae Scopoli (Homoptera: Aphididae), was compared on two plants utilized at different stages of the insect’s life cycle. Adult autumn migrants (gynoparae) are monophagous, colonizing spindle (Euonymus europaeus), whereas polyphagous summer winged aphids (alate virginoparae) are associated with a variety of herbaceous plants, including broad bean (Vicia faba). When aphids from a single clone were given access to a spindle leaf and a bean seedling in choice tests, many virginoparae settled and larviposited on both plant species over 24 h. By contrast, gynoparae showed a clear preference for spindle, with 93.5% of settled adults and 98.3% of larvae on this plant species. Close‐up video monitoring showed that gynoparae discriminated beans from spindle within a 5‐min period, whereas virginoparae behaved similarly on both plant species. For gynoparae, the major behavioural difference on the two plants appeared after a brief (epidermal) stylet penetration, with many insects taking flight within a few seconds of stylet withdrawal from bean. Factors detected during stylet insertion by gynoparae must therefore inhibit take‐off on spindle. Electrical recording experiments showed that aphids often punctured a cell membrane during brief probes on both plant species, and intracellular stylet activities always included a waveform associated with ingestion. When gynoparae puncture spindle cells their behaviour is probably modified by intracellular metabolites detected via gustation of ingested epidermal cell sap. These cues may inhibit the take‐off reflex which otherwise follows probing.     

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.     

Juvenile hormone and photoperiodically controlled polymorphism in Aphis fabae: Prenatal effects on presumptive oviparae

All three naturally occurring juvenile hormones (JH's) were shown to have effects on the parthenogenetic/gamic polymorphism of Aphis fabae; they mimicked long day conditions by inducing parthenogenetic forms. When topically applied to fourth instar gynoparae, JH caused the appearance of oviparous/viviparous intermediate morphs in the progeny. JH induced both wing development and embryogenesis in embryonic, presumptive oviparae. Embryogenesis was induced by lower doses of JH. Adult, embryo-containing alatae produced by treatment with high JH doses were capable of flight, and whilst reluctant to reproduce, their few viable progeny were oviparae. They did, however, differ from normal gynoparae in size, occasional presence of scent plaques on the metathoracic tibiae, numbers of secondary rhinaria on the antennae and morphogenetic response to postnatal rearing in long day conditions. The presumptive, oviparous embryos most sensitive to JH treatment were shown to be ca 323 μm in length, close to the stage where their germaria differentiate as parthenogenetic or gamic. Similar effects were observed in the progeny of JH-treated, teneral adult gynoparae but there was no effect on the morph of progeny of long day, alate virginoparae. The JH's differed in potency in the order JH I > JH II > JH III. The treatment of fourth instar gynoparae also induced a terminal batch of apparently normal viviparous progeny in a number of aphids. This result was obtained even at JH doses below threshold for the appearance of oviparous/viviparous intermorphs.     

各型桃蚜触角感器的比较研究

该文对桃蚜Myzus persicae(Sulzer)各型触角感器进行了比较研究。扫描电镜观察结果表明桃蚜具有钟形感器、毛形感器、原生感觉圈和次生感觉圈4种类型的触角感器。桃蚜各型触角感器的最大差异主要表现在次生感觉圈上,雄蚜、雌性母和有翅孤雌蚜具有次生感觉圈,雌性蚜、干母和无翅孤雌蚜无;但雄蚜具有的次生感觉圈数目多于雌性母和有翅孤雌蚜,且不仅仅分布在触角第3节,第4、5节也有分布。本文探讨了桃蚜触角感器在化学生态学上的功能作用,各型触角感器的差异与桃蚜寄主选择、迁移及交配行为的关系,并进一步分析了桃蚜触角感器的性二型现象。     

Wing development and parthenogenesis induced in progenies of kinoprene-treated gynoparae of Aphis fabae and Myzus persicae

Presumptive gynoparae of Aphis fabae and Myzus persicae were exposed to various levels of kinoprene (Zoecon's ZR 777) by being placed as 4th-instar alatiform larvae on bean or radish seedlings that had been sprayed with different concentrations of kinoprene in an acetone-tween-water emulsion. Larvae exposed to the highest (0.1%) concentration tested developed into adults 1 to 2 days sooner than those on control plants. The adults on the treated plants had variously deformed wings, reduced sclerotization (and pigmentation in the case of M. persicae) and other apteriform features. On reaching adulthood the affected aphids settled to feed and started to larviposit some days earlier than the control aphids. After two weeks as adults, treated gynoparae of M. persicae produced more larvae than the 7 to 9 typically deposited by control gynoparae under the short-day and cool temperature conditions employed in these tests.Whereas most or all of the larvae produced by the control gynoparae developed into oviparae (apterous, egg-laying, sexual females), gynoparae exposed to 0.1% kinoprene-treated plants predominantly produced alatiform viviparous offspring. If the latter were allowed to develop on untreated plants they deposited a few oviparous larvae. Alatiform virginoparae of M. persicae (from the same holocyclic strain that produced the gynoparae) also responded to kinoprene by developing wing deformities and by producing alatiform offspring. In contrast, alatiform virginoparae from an androcyclic strain of M. persicae, although developing wing deformities, produced only apterous progeny.The stimulation by kinoprene of wing development and parthenogenesis in the progeny of treated gynoparae is discussed in the light of our present knowledge of these aspects of aphid polymorphism.     

Production of males and gynoparae by apterous viviparae of Myzus persicae continuously exposed to different scotoperiods

At 18–19°C, apterous viviparae of a holocyclic strain of Myzus persicae raised from birth under constant scotoperiods of 9 hr 26 min–15 hr darkness per diem deposited apterous viviparae, alate viviparae, and males, in that sequence. Under the shortest scotoperiod (9 hr 26 min) in which males were recorded, only a few were deposited at the end of reproduction. With increasing duration of the scotoperiods, the aphids switched to the exclusive deposition of males progressively earlier in their reproductive lives. Thus more males and fewer females were produced; however the proportion of wingless females increased. Similar trends were recorded when the number of prenatal exposures to each scotoperiod was increased from 0 to 7.Alatae developed into gynoparae at scotoperiods of 10 hr 29 min or longer, and into virginoparae at 9 hr 40 min or shorter scotoperiods. Gynoparae and alate virginoparae, as well as alates that produced both oviparae and viviparae were found at 10 hr and 10 hr 15 min.When apterous viviparae were transferred to scotoperiods of 10 hr 29 min or 15 hr only when they attained adulthood, they also deposited males but only toward the end of their reproductive lives.     

Antibiotics, primary symbionts and wing polyphenism in three aphid species

The possible role of the primary Buchnera symbionts in wing polyphenism is examined in three aphid species. Presumptive winged aphids were fed on antibiotic-treated beans to destroy these symbionts. As previously reported, this leads to inhibited growth and low/zero fecundity. When such treatment is applied to the short-day-induced gynoparae (the winged autumn migrant) of the black bean aphid, Aphis fabae, it also causes many insects to develop as wingless or winged/wingless intermediate adult forms (apterisation). However, whilst antibiotic treatment of crowd-induced, long-day winged forms of the pea aphid, Acyrthosiphon pisum (a green and a pink clone) and the vetch aphid, Megoura viciae has similar effects on size and fecundity, it does not affect wing development. Food deprivation also promotes apterisation in A. fabae gynoparae but not in the crowd-induced winged morphs of the other two species. Thus, it appears that apterisation in A. fabae is not a direct effect of antibiotic treatment or a novel role for symbionts but is most likely related to impaired nutrition induced by the loss of the symbiont population.     

Responses of holocyclic and anholocyclic Rhopalosiphum padi populations to low‐temperature and short‐photoperiod induction

The different life cycles of aphid species make these organisms good models for studying the short‐term consequences of sex. The bird cherry‐oat aphid Rhopalosiphum padi has a wide geographic distribution and correspondingly different life cycles. In this study, the life cycles of R. padi collected from six different regions in China were characterized experimentally by comparing the responses of holocyclic and anholocyclic populations to low‐temperature and short‐photoperiod induction. Clones collected from Chuzhou, Taian, and Taigu consistently reproduced via obligate parthenogenesis, whereas clones from Hami and Baicheng were holocyclic in their response, and those from Lanzhou were both holocyclic and anholocyclic. Prolonged exposure to low temperature and a short photoperiod (LS) had negative effects on the offspring of anholocyclic aphids with regard to adult lifespan, total longevity, and fecundity compared with aphids maintained at a normal temperature and a long photoperiod (NL). Holocyclic LS R. padi had longer developmental times at all nymph stages, a shorter adult lifespan, shorter total longevity, and a lower fecundity than NL counterparts. The adult prereproduction period of gynoparae was significantly longer than that of virginoparae, and the total longevity of gynoparae was significantly shorter than that of virginoparae. Moreover, the net reproductive and gross reproduction rates, as well as the total fecundity, were roughly fivefold higher in virginoparae than in gynoparae, indicating that there is the short‐term cost of sex. When maintained on their secondary host (Triticum aestivum), gynoparae, males, and oviparae produced by holocyclic populations could survive, and gynoparae produced oviparae. However, under NL conditions, oviparae could not produce overwintering eggs on the secondary host, whereas a few overwintering eggs were generated by oviparae under LS conditions. Taken together, these results illuminate the complexity of insect responses and contribute to a complete understanding of the aphid life cycle and its evolution.     

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.     

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.