Feeding responses of two grain aphids to barley yellow dwarf virus-infected oats

The probing behavior of greenbugs, Schizaphis graminum (Rondani), and oat-bird cherry aphids, Rhopalosiphum padi (L.), was electronically monitored on healthy oats, Avena sativa (L.), and on oats infected with the RPV-NY isolate of barley yellow dwarf virus (BYDV). S. graminum fed better on infected oats than on healthy oats. This was manifested by a shorter time before initiation of committed phloem ingestion, and by increased duration of ingestion from phloem of infected compared to healthy plants. In addition, on infected oats S. graminum made fewer interruptions in their probing once their stylets were inserted into tissues, including phloem. R. padi fed similarly on infected and healthy oats, except that these aphids made fewer short probes to the phloem (lasting <15 min) on infected compared to healthy oats. BYDV infection of oats increased the rate of population growth of S. graminum relative to that on healthy oats but had no effect on the population growth of R. padi. The proportion of aphids of R. padi which developed into alatae on BYDV-infected oats was significantly greater than on healthy oats, but S. graminum showed no such response.

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Résumé Le comportement de sondage de S. graminum (Rond.) et de R. padi (L.) a été suivi électroniquement sur avoine (Avena sativa L.) saine ou contaminée par l'extrait RPV-NY du virus jaune du nanisme de l'orge (BYDV). S. graminum s'est mieux alimenté sur avoine contaminée que saine; ceci se traduisait par un temps de latence inférieur avant l'ingestion de phloème et par une prolongation de la période d'ingestion. De plus, sur avoine contaminée, S. graminum a moins souvent interrompu le sondage, une fois que les stylets avaient étè insérés dans les tissus, y compris le phloème. Les résultats avec R. padi présentaient les mêmes tendances, mais les améliorations des performances alimentaires de cette espèce sur avoine contaminée ont été moins nettes que pour S. graminum. La contamination de l'avoine par BYDV a accru le taux de croissance de la population de S. graminum par rapport à ce qui a été observé sur avoine saine; l'effet était moins prononcé pour R. padi. La proportion de pucerons devenant aliés était plus élevée sur avoine contaminée que saine, rien de tel n'a été observé avec S. graminum.

     

The role of sinigrin in host plant recognition by aphids during initial plant penetration

Plant penetration behaviour (probing) of the cabbage aphid, Brevicoryne brassicae, and the pea aphid, Acyrthosiphon pisum, was studied on excised leaves of broad beans, Vicia faba, kept in water or in a 1% aqueous solution of sinigrin. Using the DC EPG (Electrical Penetration Graph) technique it was shown that the cabbage aphid on sinigrin-untreated bean leaves showed numerous short probes into epidermis and mesophyll. None of these aphids showed either phloem salivation or ingestion waveforms on untreated leaves. In contrast, on sinigrin-treated bean leaves, 35% of the probing time was spent on phloem sap ingestion (E2) and almost all aphids reached phloem vessels and started feeding. The duration of phloem salivation before phloem ingestion and the mean duration of phloem ingestion periods were similar on a host and a sinigrin-treated non-host plant. However, the total probing time by B. brassicae was 10% longer, the total phloem sap ingestion time was twice as long, and the time to the first phloem phase within a probe was three times shorter on the host plant compared to sinigrin-treated broad beans. Acyrthosiphon pisum also responded to the addition of sinigrin to broad beans, but in this case sinigrin acted as a deterrent. On sinigrin-treated leaves, A. pisum terminated probes before ingestion from phloem vessels, and none of these aphids showed phloem salivation and ingestion on treated leaves. Glucosinolates were detected in the mesophyll cells of the brassicaceous plant, Sinapis alba. Based on this finding and in addition to the foregoing EPG analysis of aphid probing on these plants and broad beans, our hypothesis is that aphids may recognise their host plants as soon as they probe the mesophyll tissue and before they start ingestion from phloem vessels.     

Aphid feeding deterrency of ellagitannins, their phenolic hydrolysis products and related phenolic derivatives

Feeding deterrent activities of ellagic acid, two ellagitannins, gallic acid, pyrogallol, and several gallic acid derivatives towards three species of aphids were determined. The most sensitive species tested was Schizaphis graminum (Rondani), the least sensitive was Acyrthosiphon pisum (Harris). Myzus persicae (Sulzer) was of intermediate sensitivity. Ellagic acid (ED₅₀=15 ppm) and n-decyl gallate (ED₅₀=16 ppm) were particularly potent against S. graminum, while n-octyl gallate was the most active compound tested against A. pisum (ED₅₀=182 ppm) and M. persicae (ED₅₀=56 ppm). The ellagitannins, geraniin and pedunculagin, were active against S. graminum and M. persicae, but not against A. pisum. Methylation of the free hydroxyl groups of gallic acid resulted in a large decrease in activity, while esterification of its carboxyl group with alkyl chains of increasing length resulted in increasing activity against S. graminum. Against A. pisum and M. persicae, ellagic acid, gallic acid and 3,4,5-trimethoxybenzoic acid were inactive, whereas pyrogallol and the gallate esters were at least moderately active as feeding deterrents.

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Résumé L'examen a porté sur l'action répulsive, lors de l'alimentation de trois espèces de pucerons, de l'acide ellgique, de deux ellagitanins, de l'acide gallique, du pyrogallol et de plusieurs dérivés de l'acide gallique. Schizaphis graminum Rondani a été l'espèce la plus sensible, tandis que Acyrthosiphon pisum. Harris a été la moins sensible; la sensibilité de Myzus persicae Sulzer était intermédiaire. L'acide ellagic (ED₅₀=15 ppm) et le n-décyl gallate (ED₅₀=16 ppm) ont été particulièrement actifs contre S. graminum, tandis que le n-octyl gallate a été le produit le plus actif contre A. pisum (ED₅₀=182 ppm) et M. persicae (ED₅₀=56 ppm). Les ellagitanins, géraniine et pédunculagine ont été actifs contre S. graminum et M. persicae, mais pas contre A. pisum. La méthylation des groupes hydroxyl libres de l'acide gallique a réduit fortement l'activité, tandis qu l'estérification de son groupe carboxyl avec des chaînes alkyl de longuers croissantes a augmenté l'activité contre S. graminum. Les acides ellagique, gallique et 3,4,5-triméthoxybenzoïque ont été inactifs contre A. pisum et M. persicae, tandis que le pyrogallol et les esters de gallate on été pour le moins des répulsifs modérément actifs au cours de l'alimentation.

     

Effect of cultivars with varying levels of resistance to aphids on development time,sex ratio,size and longevity of the parasitoid <Emphasis Type="Italic">Aphidius colemani</Emphasis>

The influence of cultivars of common cabbage, Brassica oleracea var. capitata with varying levels of resistance to Brevicoryne brassicae (L.) and Myzus persicae (Sulzer) on key biological characteristics of Aphidius colemani (Viereck) was investigated under laboratory conditions. The total development time for female parasitoids reared on M. persicae did not differ significantly between Minicole (green-leaved, partially resistant with antibiosis factors for B. brassicae) and Derby Day (green-leaved, susceptible to both aphid species); but development was significantly faster (ca 10%) on Ruby Ball (red-leaved, partially resistant with antixenosis factors for B. brassicae). Total development time for females reared on B. brassicae was slightly shorter on Ruby Ball than on Minicole. Males reared on M. persicae developed into adults significantly faster (ca 10%) on Ruby Ball than on Minicole. However, when B. brassicae was the host, no significant variations in development time were observed. Sex ratios, size and longevity of both male and female parasitoids on either host were not significantly influenced by cultivar. The results are discussed in relation to the compatible utilisation of host-plant resistance and biological control in the integrated management of aphids.     

Effects of the antifeedant polygodial on plant penetration by aphids, assessed by video and electrical recording

Adult apterousMyzus persicae (Sulz.) discriminated within 2 min between mature Chinese cabbage (Brassica pekinensis L.) leaf halves treated with (±)-polygodial solution and solvent alone, and walked off leaf areas treated with polygodial faster than off the solvent-treated areas. However, when aphids were attached to a fine gold wire and stylet penetration of cabbage leaves was recorded electrically, polygodial treatment did not affect the number or duration of electrically-recorded penetrations, time taken to initiate a first penetration, or total penetration time. As the tethering of aphids causes behavioural restrictions which may negate the response to polygodial, indications of stylet penetration by freely-moving insects were sought. Simultaneous electrical recording and video monitoring showed that stylet penetration duration could be accurately inferred from antennal and body movements, enabling assessment of penetration times without attachment to the wire tether. When freely-movingM. persicae were video recorded during 15 min access to cabbage seedlings, polygodial treatment again had no apparent effect on stylet penetration. However, when aphids were presented with a choice of polygodial- and solvent-treated sides of floating mature cabbage leaf discs, video recordings revealed that the insects spent less time and made fewer penetrations on the polygodial-treated side. In addition to this rapid repellent effect, prolonged exposure to polygodial also produced behavioural changes. After being held for 24 h on polygodial-treated leaves or green paper prior to behavioural examination, aphids penetrated seedlings fewer times but for longer periods. The relevance of the results to virus transmission studies is discussed.     

Effects of dietary sucrose concentration on aphid honeydew carbohydrate levels and rates of excretion

Honeydew droplets freshly excreted by two aphid species, Myzus persicae (Sulz.) and Acyrthosiphon pisum (Harr.), feeding on nutritionally adequate, artificial diets containing 10–40% w/v sucrose, were collected in oil. The volume, frequency of excretion, and carbohydrate content of the droplets were determined by measuring the diameter of the droplets in the oil, by recording the number of droplets excreted per aphid per hour, and by using a quantitative colorimetric method for estimating carbohydrates. Rates for the intake or excretion of fluid, the intake and absorption of sucrose, and the excretion of carbohydrate were consequently calculated for the two species on each dietary sucrose concentration. The percentage of dietary sucrose absorbed by the aphids ranged from 36% on diet with 40% sucrose to 97% on diet with 10% sucrose. Such honeydew/diet relationships may allow one to use honeydew carbohydrate concentrations to estimate the sucrose concentrations of the phloem sap ingested by aphids feeding on host plants.

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Résumé Des gouttelettes de miellat fraîchement excrétées par deux espèces de pucerons, Myzus persicae (Sulz.) et Acyrthosiphon pisum (Harr.), pucerons nourris sur des régimes nutritifs synthétiques contenant 10–40% de saccharose, ont été récupérées dans de l'huile. Le volume des gouttelettes a été déterminé en mesurant leur diamètre individuel dans l'huile, la fréquence d'excrétion établie par le nombre de gouttelettes excrétées par puceron par heure, et la teneur en glucides par une méthode d'analyse colorimétrique quantitative. Les taux d'ingestion et d'excrétion de liquide, d'ingestion et d'absorption de saccharose ainsi que les taux d'excrétion de glucides ont été par la suite calculés pour les deux espèces d'aphides nourris sur les différentes concentrations de saccharose. Le pourcentage de saccharose alimentaire absorbé par les aphides a été de 36% sur le régime contenant 40% de ce glucide, mais s'est élevé à 97% sur le régime ne contenant que 10% de saccharose. D'après ces relations miellat/régime nutritif, et en connaissant la teneur du miellat en glucides, on peut faire une approximation de la concentration du saccharose dans la sève élaborée du phloème ingérée par les aphides.

     

Genetic variation in asexual populations of two aphids in the genus Acyrthosiphon, from an Australian lucerne field

Twelve lines of Acyrthosiphon kondoi Shinji and Acyrthosiphon pisum (Harris) were sampled from a small lucerne field in Australia and tested to assess genetic variation by measuring their responses to three lucerne cultivars. Seven or eight life history traits were assessed for each species. Differences in the responses of these traits to the cultivars revealed that three to seven genetically distinct clones were represented in the samples of 12 lines for each species. We conclude that the populations of these two aphids are variable in Australia, although they were recently introduced and reproduce asexually. The responses of the aphids to the three cultivars are described. Because of genetic variation in host plant responses of the aphids, cultivars with potential aphid resistance must be exposed to a large number of clones to confirm the presence of useful aphid resistance.

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Résumé Douze lignées d'Acyrthosiphon kondoi Shinji et d'A. pisum Harris ont été échantillonnées dans un petit champ de luzerne australien et leur variabilité génétique examinée en comparant les réponses à 3 cultivars de luzerne. De 7 à 8 caractères biologiques avaient été retenus pour chaque espèce. Les différences observées entre les réponses suivant la distribution de ces caractères chez les pucerons, ont montré que 3 à 7 clones génétiquement distincts étaient représentés dans les 12 lignées de chaque espèce. Nous en concluons que les populations de ces deux espèces de pucerons sont polymorphes en Australie, quoiqu'elles se multiplient d'une façon asexuée et aient été récemment introduites. Les réactions des pucerons aux 3 cultivars sont indiquées. Par suite de la variabilité génétique des réactions des plantes hôtes aux pucerons, des cultivars à résistance potentielle élevée devraient être exposés à un grand nombre de clones de pucerons pour confirmer la présence d'une résistance utilisable.

     

Divergent responses of two cereal aphids to previous infestation of their host plant

The role of pollen odour in resource location by the pollen beetle, Meligethes aeneus (Fabricius) (Coleoptera: Nitidulidae), a pollen-feeding insect regarded as a pest of oilseed rape, Brassica napus L., (Brassicaceae) crops, was investigated in a linear track olfactometer. Both male and female beetles were attracted to the odour of whole oilseed rape flowers, indicating that these insects can locate their host plants using floral odours as cues. The attractive odour of flowers was found to emanate from all floral parts tested: the petals/sepals, the anthers, and from pollen itself. Therefore, at least part of the attractive odour of oilseed rape flowers emanates from pollen. Beetles were more attracted to floral samples containing anthers than those without anthers when these odours were directly compared in a choice-test, and this indicates that there were detectable differences between them. Anthers and pollen may therefore release distinctive odours that are quantitatively and/or qualitatively different from the odour of the rest of the flower. These experiments support the hypothesis that pollen-seeking insects use pollen odour cues to locate this food source.     

Inducing the sexual forms and hatching the eggs of pea aphids

In temperate climates, pea aphids (Acyrthosiphon pisum) produce a single sexual generation each year in response to declining photoperiod and temperature. Mating occurs in the fall and the eggs have an obligatory winter diapause. Genetic recombination during the sexual phase is thought to be an important source of genetic variability within cyclically parthenogenetic aphid populations. Methods for reliably producing sexual forms and hatching the eggs of aphids are therefore central not only to the study of evolutionary change in aphid populations, but also for a general understanding of the origin of agriculturally important variation in destructiveness within pest species.Here, sexual forms of six pea aphid clones were induced in the laboratory and eggs were successfully hatched by creating conditions that closely mimicked those found in field situations. A declining photoperiod was produced by controlling artificial lighting using a timer with variable cycle length. Using these conditions, sexual forms were successfully produced for all six clones tested, which were then mated in all combinations. Eggs were exposed to a daily cycle of freezing and thawing in an incubator under a short-day photoperiod. Egg hatch averaged 60%, but was as high as 89% for some crosses. These methods will permit testing of evolutionary hypotheses and execution of detailed genetic studies of sources of variability within pea aphid populations. They are thus important tools for both evolutionary and agricultural studies.     

Effects of bacterial secondary symbionts on host plant use in pea aphids

Aphids possess several facultative bacterial symbionts that have important effects on their hosts'' biology. These have been most closely studied in the pea aphid (Acyrthosiphon pisum), a species that feeds on multiple host plants. Whether secondary symbionts influence host plant utilization is unclear. We report the fitness consequences of introducing different strains of the symbiont Hamiltonella defensa into three aphid clones collected on Lathyrus pratensis that naturally lack symbionts, and of removing symbionts from 20 natural aphid–bacterial associations. Infection decreased fitness on Lathyrus but not on Vicia faba, a plant on which most pea aphids readily feed. This may explain the unusually low prevalence of symbionts in aphids collected on Lathyrus. There was no effect of presence of symbiont on performance of the aphids on the host plants of the clones from which the H. defensa strains were isolated. Removing the symbiont from natural aphid–bacterial associations led to an average approximate 20 per cent reduction in fecundity, both on the natural host plant and on V. faba, suggesting general rather than plant-species-specific effects of the symbiont. Throughout, we find significant genetic variation among aphid clones. The results provide no evidence that secondary symbionts have a major direct role in facilitating aphid utilization of particular host plant species.     

Density-dependent regulation in populations of potato-colonizing aphids

Scarcity of long-term (over 30 years) data series represents a major challenge for an accurate estimation of the role of density-dependent processes in population regulation. We analyzed population densities of the wingless parthenogenic morphs of buckthorn aphid (BA), Aphis nasturtii Kaltenbach, potato aphid (PA), Macrosiphum euphorbiae (Thomas), and green peach aphid (GPA), Myzus persicae (Sulzer) from 1949 to 2003 for signs of density-dependent regulation. We found strong evidence of density-dependent regulation, with detection of density dependence being fairly consistent among the different statistical techniques. Direct density dependence was detected for the populations of all three species. There was also evidence of delayed density dependence for PA. The periodicity of population fluctuations for BA and GPA was 6.1 years and 3.9 years, respectively. The periodicity for PA was not explicit, being highly variable throughout the time series. Effects of density-independent weather factors were relatively minor compared to density-dependent regulation. The BA populations experienced a significant reduction in both density and annual oscillations starting in 1995, while GPA populations experienced a similar reduction in 1991. No such change was apparent for PA. The most likely explanation for the observed phenomenon is a change in the composition of the lady beetle community following the establishment of two alien coccinellid species, and/or changes in insecticide use by commercial growers in the area of the study.     

Strong parasitoid-mediated selection in experimental populations of aphids

Clonal diversity in asexual populations may be maintained if different clones are favoured under different environmental conditions. For aphids, parasitoids are an important variable of the biotic environment. To test whether parasitoids can mediate selection among host clones, we used experimental populations consisting of 10 clones of the peach-potato aphid, Myzus persicae, and allowed them to evolve for several generations either without parasitoids or in the presence of two species of parasitoid wasps. In the absence of parasitoids, strong shifts in clonal frequencies occurred, mostly in favour of clones with high rates of increase. The parasitoid Diaeretiella rapae hardly affected aphid densities but changed the outcome of competition by favouring one entirely resistant clone and disfavouring a highly susceptible clone. Aphidius colemani, the more infective parasitoid, strongly reduced aphid densities and dramatically changed host clonal frequencies. The most resistant clone, not a successful clone without parasitoids, became totally dominant. These results highlight the potential of temporal or spatial variation in parasitoid densities to maintain clonal diversity in their aphid hosts.     

Sex pheromone discrimination by male aphids of a biotype of Schizaphis graminum

The ability of male aphids of two different biotypes (C & E) of the greenbug, Schizaphis graminum (Rondani), to distinguish ovipara-produced sex pheromone of their own biotype from that of the other biotype was examined using an arena olfactometer. Biotype E males showed a strong preference for biotype E oviparae; the preference of biotype C males for biotype C oviparae was less marked. These behavioral findings indicate a potential biochemical reproductive isolating mechanism for these biotypes.

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Unterschiedung der weiblichen geschlechtspheromone zweier biotypen der aphide Schizaphis graminum durch die männchen

Zusammenfassung Die Reaktion der Männchen von zwei Biotypen (C & E) der Aphide Schizaphis graminum auf die Geschlechtspheromone der Oviparen ihres eigenen und des anderen Biotyps wurde mit Hilfe einer Olfaktometer-Arena untersucht. Biotyp E Männchen zeigten eine starke Präferenz für Biotyp E Ovipare, während die Präferenz von Biotyp C Männchen für Biotyp C Ovipare weniger stark ausgeprägt war. Diese Resultate deuten auf einen möglichen verhaltensbiologischen Isolationsmechanismus der beiden Biotypen dieser Blattlausart in der Natur hin.

     

Polymer webs to prevent virus transmission by aphids in seed potatoes

Electrical registration and visual observation of penetration behaviour combined with ELISA-technique showed that laboratory-made webs of synthetic fibers were very promising to protect plants against virus infection by aphids. It was demonstrated that stylet penetration by aphids into potato leaves could be prevented by covering plants with these materials. The recent mass production of polypropylene sealed fiber fleeces and polyethylene nettings have made field experiments possible to determine their effect on aphid probing behaviour and virus transmission.Field experiments with potatoes showed that one particular fleece completely protected the plants against virus transmission (PVY and PLRV) and with a second fleece and a netting a small proportion of plants were infected. In uncovered plots up to 25% of plants became infected. A seed potato crop covered with polypropylene fleece which is inspected regularly for damage will be fully protected against virus transmission.

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Résumé Des enregistrements électriques et des observations visuelles du comportement de pénétration, combinés avec l'usage de la technique ELISA, ont montré que l'usage de toiles de fibres synthétiques élaborées au laboratoire était très prometteur pour la protection des plantes contre des infections virales transmises par des aphides. On a pu démontrer que la pénétration du stylet dans des feuilles de pomme de terre pouvait être empèchée en couvrant les plantes avec ces matériaux. La production en masse récente de nappes en fibres de polypropylene soudées et de filets de polyethylène a rendu possibles des expériences de terrain pour déterminer leur effet sur le comportement de sondage des pucerons et sur la transmission des virus.Des essais en plein champ avec des pommes de terre ont montré qu'un type de nappe en particulier protégeait complètement les plantes contre la transmission de virus (PVY en PLRV) et qu'avec un second type de nappe et un filet une petite proportion de plantes était infectée. Dans des parcelles non couvertes la proportion de plantes infectées s'élevait jusqu'à 25%. Une récolte de pommes de terre semencières couverte d'une nappe de polypropylène régulièrement inspectée afin de remédier aux détériorations éventuelles sera totalement protégée contre la transmission de virus.

     

Seasonal variation in the photoperiodic responses of a pea aphid population: evidence for long-distance movements between populations

Summary The purpose of the study was to quantify long distance movements in populations of pea aphid, Acyrthosiphon pisum (Harris), by estimating origins and distances travelled by immigrants into a southern Manitoba population. A strong relationship was demonstrated between latitude of origin and photoperiods at which pea aphid populations are stimulated to produce the diapause stage (Smith 1987). Therefore, the approach was to use photoperiodic response as a physiological marker to identify the source of immigrant aphids. The responses of 89 clones from Glenlea, Manitoba (49°38N), sampled 5 times over 2 seasons, were measured. One sample of clones collected the first season had photoperiodic responses similar to those of a population about 300 km to the south, and significantly different from clones collected in spring of the same year at the same site. Weather analysis corroberates that the migrants were probably carried into Manitoba on a southerly flow of air during the previous 24 to 36 h.     

Amplification and methylation of an esterase gene associated with insecticide-resistance in greenbugs, Schizaphis graminum (Rondani) (Homoptera: Aphididae)

The greenbug aphid, Schizaphis graminum (Rondani) has developed resistance to organophosphorus insecticides by the over-production of esterases that have been classified as Type I and Type II. The first twenty N-terminal amino acids of the Type I esterase were determined and used to design an oligonucleotide, which in conjunction with an active site primer derived from conserved sequences of other insect esterases and two internal primers specific for esterases from another aphid species resulted in a 0.85 kb genomic DNA fragment from resistant greenbugs. This was extended by 5′ RACE which provided approximately 1.2 kb of the 5′ end of the esterase gene. The 5′ DNA sequence corresponded to 19 of the 20 known amino acids of the Type I esterase, with the last needing only a one base change (probably resulting from a PCR artifact). Furthermore, the sequence showed very close similarity to the amplified E4/FE4 esterase genes of Myzus persicae (Sulzer). A comparison of sequences suggested that the S. graminum gene has introns in the same positions as the first two introns of E4/FE4, with the second intron being considerably larger in S. graminum. Probing of Southern blots with the 0.85 kb esterase fragment showed that the gene encoding the Type I esterase is amplified 4- to 8-fold in resistant S. graminum and that the amplified sequences contain 5-methylcytosine at MspI/HpaII sites, again in agreement with previous findings for M. persicae genes.