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.     

Effects of wing polyphenism, aphid genotype and host plant chemistry on energy metabolism of the grain aphid, Sitobion avenae

Wing dimorphism has been proposed as a strategy to face trade-offs between flight capability and fecundity. In aphids, individuals with functional wings have slower development and lower fecundity compared with wingless individuals. However, differential maintenance costs between winged and wingless aphids have not been deeply investigated. In the current study, we studied the combined effect of wing dimorphism with the effects of aphid genotypes and of wheat hosts having different levels of chemical defences (hydroxamic acids, Hx) on adult body mass and standard metabolic rates (SMR) of winged and wingless morphs of the grain aphid, Sitobion avenae. We found that wingless aphids had higher body mass than winged aphids and that body mass also increased towards host with high Hx levels. Furthermore, winged aphids showed a plastic SMR in terms of Hx levels, whereas wingless aphids displayed a rigid reaction norm (significant interaction between morph condition and wheat host). These findings suggest that winged aphids have reduced adult size compared to wingless aphids, likely due to costs associated to the development of flight structure in early-life stages. These costs contrast with the absence of detectable metabolic costs related to fuelling and maintenance of the flight apparatus in adults.     

Differential regulations of wing and ovarian development and heterochronic changes of embryogenesis between morphs in wing polyphenism of the vetch aphid

SUMMARY In wing polyphenisms that produced alternative wing morphs depending on environmental conditions, the developmental regulations to balance between flight and reproductive abilities should be important. Many species of aphids exhibit wing polyphenisms, and the development of wing and flight muscles is thought to incur costs of reproductive ability. To evaluate the relationship between flight and reproduction, the fecundity and the wing- and ovarian development in the parthenogenetic generations were compared between winged and wingless aphids in the vetch aphid Megoura crassicauda . Although no differences in offspring number and size were detected, the onset of larviposition after imaginal molt was delayed in winged adults. The comparison of growth in flight apparatus revealed that, after the second-instar nymphs, the flight-apparatus primordia of presumptive wingless aphids were degenerated while those of winged nymphs rapidly developed. In the ovaries of winged line, the embryo size was smaller and the embryonic stages were delayed from third to fifth instars, although these differences had disappeared by the time of larviposition. It is therefore likely that the delay in larviposition in winged aphids is due to the slower embryonic development. The correlation between embryo size and developmental stage suggests that the embryos of winged aphids are better developed than similarly sized embryos in wingless aphids. These heterochronic shifts would facilitate the rapid onset of larviposition after the dispersal flight. This developmental regulation of embryogenesis in the aphid wing polyphenism is suggested to be an adaptation that compensates the delay of reproduction caused by the wing development.     

The cost of being able to fly in the milkweed-oleander aphid,Aphis nerii (Homoptera: Aphididae)

Summary In the wing dimorphic milkweed-oleander aphid,Aphis nerii, winged aphids begin reproducing about 1.5 days after wingless aphids. The longer maturation period is primarily due to slower development since even adult eclosion by winged aphids takes place after wingless aphids begin reproducing. The delay is not due to a post-eclosion, pre-reproductive flight since, beginning with the fourth instar, larval winged aphids were reared at a density of one per plant and the vast majority were not stimulated to fly under such low-density conditions. Thus, the ability to fly incurs a fitness cost in terms of delayed reproduction, irrespective of whether flight actually occurs. We did not observe a difference between morphs for lifetime fecundity, even though wingless aphids have larger abdomens than winged aphids and for both morphs there is a significant correlation between abdomen width and fecundity. Offspring produced by wingless aphids over the first four days of reproduction are larger than those produced by winged aphids, and the size difference at birth is maintained into adulthood. However, there are no differences in life history traits between these offspring, including maturation period and lifetime fecundity. Thus, reduced body size does not increase the cost of being able to fly, at least under the conditions of these experiments. The cost of being able to fly in this species should favor reduced production of winged individuals in populations that exploit more permanent host plants.     

The visual responses of free-flying summer and autumn forms of the black bean aphid, Aphis fabae, in an automated flight chamber

ABSTRACT Summer (virginoparae) and autumn (gynoparae) winged forms of Aphis fabae were flown in an automated flight chamber for up to 100 min, while being exposed to a small green target at their own level for 5 s of each minute. Virginoparae climbed more slowly than gynoparae and responded to the target by turning towards it throughout experimental flights. Gynoparae, on the other hand, were almost completely unresponsive to the target, although some individuals began to respond after an hour or more of flight. The differences between the two migratory forms are discussed with reference to previous work on aphid flight behaviour in the field and the laboratory.     

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.     

The influence of natural enemies on wing induction in Aphis fabae and Megoura viciae (Hemiptera: Aphididae)

Previous studies have shown that the aphid species, Aphis fabae Scopoli and Megoura viciae Buckton, do not produce winged offspring in the presence of natural enemies, in contrast to results for the pea aphid (Acyrthosiphon pisum (Harris)) and the cotton aphid (Aphis gossypii Glover); but these studies did not involve exposing aphids directly to natural enemies. We exposed colonies of both A. fabae and M. viciae to foraging lacewing (Chrysoperla carnea (Stephens)) larvae and found that the predators did not induce winged morphs among offspring compared to unexposed controls. Colonies of A. fabae responded to an increase in aphid density with increasing winged morph production, while such response was not found for M. viciae. We suggest that different aphid species differ in their susceptibility to natural enemy attack, as well as in their sensitivity to contact.     

Host acceptance by aphids: probing and larviposition behaviour of the bird cherry-oat aphid, Rhopalosiphum padi on host and non-host plants

The probing and larviposition behaviour of the bird cherry-oat aphid, Rhopalosiphum padi on summer and winter host plants were investigated using electrical penetration graph (EPG) coupled with simultaneous video recording. In this way the precise probing history prior to parturition can be monitored and the location of possible reproductive stimulants identified. On the host plant, all gynoparae (autumn winged females that give birth to sexual females on bird cherry, Prunus padus, the primary host) and 55% of winged virginoparae (summer females which produce further virginoparae on barley, Hordeum vulgare, a secondary host) initiated larviposition before phloem contact. However, 90% of wingless virginoparae (on barley) contacted the phloem before first larviposition whilst 10% did not. Thus, phloem contact does not appear to be a pre-requisite for these aphid forms to initiate reproduction.     

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.     

Wing development in parasitized male and female Sitobion fragariae

Abstract.Different stages of presumptive winged morphs (males, gynoparae and alate virginoparae) of the blackberry‐cereal aphid, Sitobion fragariae , were exposed to attack by Aphidius ervi . Even though the mechanism influencing wing development in the three aphid morphs differs, the effects of parasitism were similar. Alatiform structures were completely inhibited in all three morphs when the initial attack took place in their first or early second stadium. The disruption of wing development also resulted in apterous/alate‐intermediate forms when aphids were attacked from first (males and gynoparae) or early second (alate virginoparae) up to the fourth larval stadium. The fact that wing development was still disrupted when aphids with well developed wingbuds were parasitized indicates that the early stages of parasitization were influential. Thus, the morphogenetic effects may be exerted by the parasitoid egg or calyx fluid.     

Maternal reproductive decisions are independent of feeding in the black bean aphid,Aphis fabae

Aphids are phloem feeders and an important assumption has been that reproduction is initiated only after phloem ingestion. Here we investigate the plant tissue location of parturition cues in winged and wingless, summer virginoparae and autumn migrants (gynoparae) of the black bean aphid, Aphis fabae. These seasonal forms have different host preferences. Using electrical penetration graph (EPG: to observe activity of the mouthparts) and video-monitoring procedures we demonstrate that the time to first parturition after host-plant contact is significantly shorter than the time to first registered phloem contact in the summer winged form. In gynoparae, the time to first parturition does not significantly differ from time to first phloem contact but is shorter than time to first phloem ingestion. Times to first parturition, first registered phloem contact and first phloem ingestion do not differ significantly in the summer wingless form. Simultaneous EPG and video recording procedures show that a high proportion of individuals of all morphs (45-70%) initiate reproduction before sustained phloem activities (salivation/ingestion). The only behaviours that all individuals demonstrate before first parturition are ‘non-penetration’ (aphid on plant surface with mouthparts outside plant) and stylet ‘pathway activity’, including secretion of gelling saliva and penetration of the non-vascular (epidermis and mesophyll) cells. A short period of penetration of the peripheral plant tissues (five cell punctures per individual) by aphids tethered and monitored by EPG decreases the time to first parturition of the winged summer form when subsequently placed on a Parafilm sachet containing 15% sucrose solution. This treatment also significantly increases the incidence of reproduction and individual reproductive output of gynoparae over a 24-h period. No detectable effects of tissue penetration on subsequent reproductive output are observed in the wingless summer form. Additionally, EPGs reveal that a number of aphids of all morphs display xylem ingestion, which occurs predominantly before initiation of phloem feeding but is not necessary to initiate parturition. It is concluded that aphids are likely to detect parturition cues during stylet punctures of cells within peripheral tissue layers, before reaching the phloem vessels.     

Genetic variation for an aphid wing polyphenism is genetically linked to a naturally occurring wing polymorphism

Many polyphenisms are examples of adaptive phenotypic plasticity where a single genotype produces distinct phenotypes in response to environmental cues. Such alternative phenotypes occur as winged and wingless parthenogenetic females in the pea aphid (Acyrthosiphon pisum). However, the proportion of winged females produced in response to a given environmental cue varies between clonal genotypes. Winged and wingless phenotypes also occur in males of the sexual generation. In contrast to parthenogenetic females, wing production in males is environmentally insensitive and controlled by the sex-linked, biallelic locus, aphicarus (api). Hence, environmental or genetic cues induce development of winged and wingless phenotypes at different stages of the pea aphid life cycle. We have tested whether allelic variation at the api locus explains genetic variation in the propensity to produce winged females. We assayed clones from an F2 cross that were heterozygous or homozygous for alternative api alleles for their propensity to produce winged offspring. We found that clones with different api genotypes differed in their propensity to produce winged offspring. The results indicate genetic linkage of factors controlling the female wing polyphenism and male wing polymorphism. This finding is consistent with the hypothesis that genotype by environment interaction at the api locus explains genetic variation in the environmentally cued wing polyphenism.     

Visual responses of flying aphids and their chemical modification

Abstract. Five winged forms of the black bean aphid, Aphis fabae, were subjected to 30 min of free flight in a recently designed, automated wind tunnel (David & Hardie, 1988). A circular green target, on the side of the chamber, was illuminated for 2.5 s of each minute of flight. Half of the fundatrigeniae ('spring migrants') and all of the virginoparous 'summer migrants' responded by flying towards the illuminated target while gynoparae ('autumn migrants') induced by short days over two generations in uncrowded conditions were unresponsive. 27% of generation 1 gynoparae, induced by crowded conditions and short days over one generation, responded to the target as did 50% of males.
Anaesthetization of adult gynoparae with nitrogen and carbon dioxide, but not ether, induced targeted flights. However, the effect of carbon dioxide was transient over the first 10 min whereas the effect of nitrogen continued throughout the 30-min experimental period. In addition, treatment of late fourth instar larvae, but not young adults, with Juvenile Hormone I provoked target responses. The results are discussed with reference to settling behaviour and the endocrine control of polyphenism.     

Differential photoperiodic responses in genetically identical winged and wingless pea aphids, Acyrthosiphon pisum, and the effect of day length on wing development

Abstract. Winged and wingless individuals of a pink clone of the pea aphid, Acyrthosiphon pisum (Harris), showed differences in the response curves for photoperiodic induction of both males and sexual females (oviparae). The critical night length (CNL) for ovipara induction in winged aphids was 0.75 h shorter than in wingless aphids, whereas the CNL for male induction in winged aphids was 1.0h longer than in wingless aphids. This means that in winged aphids the CNL for male induction in winged aphids was 0.5 h longer than that for ovipara induction, while in wingless aphids the CNL for male induction was 1.0–1.5 h shorter than that for ovipara induction, and also the shapes of the curves differed.
Winged aphids were produced by wingless mothers which were crowded as young adults. However, when young adults were crowded in long nights, winged aphids were not produced, and the CNL for wing inhibition was between 9.5 and 10h. This effect of photoperiod on wing induction was maternal.     

禾谷缢管蚜复眼外部形态的扫描电镜观察

【目的】通过比较禾谷缢管蚜Rhopalosiphum padi两种生活史(全周期型和不全周期型)及5种蚜型(有翅孤雌蚜、无翅孤雌蚜、雄蚜、雌性蚜和产雌性母)成蚜复眼外部形态,分析了其视觉能力差异,为进一步探索禾谷缢管蚜生殖转换及寄主搜寻机制提供参考。【方法】利用扫描电镜技术,观察全周期型和不全周期型禾谷缢管蚜无翅孤雌蚜及其低温短日照(12℃, 光周期8L∶16D)诱导条件下全周期型所产5种蚜型成蚜的复眼外部形态。【结果】禾谷缢管蚜成蚜复眼着生于头部触角基部,呈卵圆形,小眼间无感觉毛。全周期型和不全周期型禾谷缢管蚜无翅孤雌蚜的复眼大小(长轴直径及短轴直径)、复眼弧高、小眼数量和小眼直径均没有显著差异。而这5个复眼指标在低温短日照诱导全周期型禾谷缢管蚜产生的5种蚜型的成蚜中均存在显著性差异,其中,雄蚜、有翅孤雌蚜和产雌性母的复眼区域较大,小眼数量较多;雌性蚜的复眼区域最小,小眼数量也最少。【结论】综合以上指标结果,推测5种蚜型的视觉能力由大到小是：雄蚜＞有翅孤雌蚜＞产雌性母＞无翅孤雌蚜＞雌性蚜。复眼和小眼的这些差异可能与禾谷缢管蚜生殖转换及寄主搜寻存在一定的联系。     

Competition and dispersal in the pea aphid: clonal variation and correlations across traits

Abstract.  1. The presence of an across-species trade-off between dispersal ability and competitive ability has been proposed as a mechanism that facilitates coexistence. It is not clear if a similar trade-off exists within species. Such a trade-off would constrain the evolution of either trait and, given appropriate selection pressures, promote local adaptation in these traits.
2. This study found substantial levels of heritable variation in competitive ability of the pea aphid, Acyrthosiphon pisum Harris (Homoptera: Aphididae), measured in terms of relative survival when reared with a single clone of the vetch aphid, Megoura viciae Buckton (Homoptera: Aphididae).
3. Pea aphids can move to new patches by either flying (longer distance dispersal) or walking (local dispersal) from plant to plant. There was considerable clonal variation in dispersal ability, measured in terms of the proportion of winged offspring produced, and ability to survive away from their host plant.
4. Winged individuals showed longer off-plant survival times than wingless forms of the same pea aphid clone.
5. There was no evidence of a relationship between clonal competitive ability and either measure of dispersal ability, although the power of the test is limited by the number of pea aphid clones used in the trial.
6. However, there was a positive correlation between clonal fecundity and the proportion of winged offspring produced. Although speculative, it is suggested that clones that are more likely to either overwhelm their host plant or attract higher numbers of natural enemies as a result of having higher fecundity are more likely to produce winged morphs.     

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.     

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.     

Genotypic variation among different phenotypes within aphid clones.

Most aphid species Hemiptera: Aphididae are parthenogenetic between periods of sexual reproduction. They are also highly polyphenic, with different adult morphs occurring in the life cycle, piz. winged, wingless, asexual and sexual. It is assumed that aphids born in a parthenogenetic clonal lineage are genetically identical regardless of the final adult form with the exception of sexual forms). Using the randomly amplified polymorphic DNA-polymerase chain reaction (RAPD-PCR) we have found that different asexual adult phenotypes winged and wingless of some clones of two cereal aphid species (the grain aphid, Sitobion avenae (F.) and the bird-cherry aphid. Rhopalosiphum padi (L.) may be distinguished by the presence or absence of one or more RAPD-PCR bands. In three of nine clones examined, such differences were found, and Southern blotting and hybridization of the discriminating bands confirmed these to be of aphid origin, rather than due to endosymbiotic bacteria or contaminating fungi. The main 248 and 296 bp bands, in the two species respectively, were sequenced and found to be A/T rich. The smaller band showed 57% homology with white striated muscle over a stretch of 90 bp. Genomic DNA treated with dimethyl sulphoxide to remove secondary structures still showed differences in RAPD-PCR profiles between winged and wingless morphs within the unusual clones. This discovery may be widespread and therefore it is important to understand the phenomenon in relation to clonal organisms.