Electronically monitored cowpea aphid feeding behavior on resistant and susceptible lupins

The feeding behavior of cowpea aphid, Aphis craccivora Koch (Homoptera: Aphididae) was examined on seedlings of narrow leafed lupin, Lupinus angustifolius L., and yellow lupin, L. luteus L., using electronic monitoring of insect feeding behavior (EMIF). Aphid feeding behavior was first compared between resistant (cv. Kalya) and susceptible (cv. Tallerack) varieties of narrow-leafed lupin. Aphids spent significantly more time in non- penetration and stylet pathway activities, and significantly less time in the sieve element phase on Kalya than on Tallerack, suggesting that feeding deterrence is an important component of aphid resistance in Kalya. Aphid feeding on a susceptible yellow lupin variety (cv. Wodjil) was then compared with that on two resistant lines, one (Teo) with high and the other (94D024-1) with low seed alkaloid content. There were no consistent differences in aphid feeding behavior between Wodjil and Teo. Total, mean and percentage sieve element phase times were significantly lower, and total and percentage times in non-phloem phase were greater on 94D024-1 than on Wodjil, suggesting the possibility of phloem-based deterrence in 94D024-1.     

Cowpea aphid performance and behaviour on two resistant cowpea lines

In a series of laboratory and climate chamber tests we compared the growth and behaviour of Aphis craccivora on one susceptible (ICV-1) and two aphid-resistant (ICV-11 and ICV-12) cowpea lines. The aphids' growth rates were much lower on the resistant cowpea lines than on the susceptible one, indicating strong antibiosis. In addition, the aphids invariably settled in higher numbers on the susceptible line than on either of the resistant. Compared to ICV-1, damaged leaves of the resistant line ICV-12 were settled upon to a higher degree than undamaged leaves, and leaf discs from the same line were even less resistant.On resistant lines individual aphids waited a significantly longer time before making their first test probe. Total probing time as well as the time preceding a decision to stay or leave was also longer.These results are discussed in relation to the possible mechanisms involved, and we also consider the effects of previous leaf feeding on the expression of resistance in the field.

---

Résumé Au cours d'expériences au laboratoire et en chambres climatisées nous avons comparé la croissance et le comportement de A. craccivora sur une lignée sensible (ICV-1) et deux lignées résistantes (ICV-11 et ICV-12) de V. unguiculata. Les vitesses de croissance des pucerons ont été beaucoup plus lentes sur les lignées résistantes que sur la lignée sensible, ce qui révèle une forte antibiose. De plus, les pucerons atterrissent invariablement en plus grand nombre sur la variété sensible. Par comparaison avec ICV-1, les atterrissages sur lignée résistante ICV-12 étaient plus nombreux sur les feuilles endommagées que sur les feuilles intactes; les disques de feuilles de cette même lignée étaient encore moins résistants.Les pucerons ont séjourné individuellement un temps plus long sur les lignées résistantes avant de faire leur premier sondage. Le temps consacré aux sondages ainsi que le temps précédant de choix entre départ ou maintien sur la feuille étaient plus longs avec les lignées résistantes.Ces résultats ont été discutés en fonction des mécanismes impliqués. Nous avons aussi examiné les effets de la consommation antérieure sur les manifestations de la résistance dans la nature.

     

Predation byCheilomenes vicina [Coleoptera: Coccinellidae] on the cowpea aphid,Aphis craccivora [Homoptera: Aphididae]: Effect of prey stage and density

Consumption of larvae and females ofAphis craccivora Koch by 1st and 4th larvae and adults ofCheilomenes vicina (Muls.) was studied under fluctuating temperature (24–30°C). The early aphid instars were consumed in significantly greater numbers than later instars and females. The feeding rates ofC. vicina were significantly positively correlated with the population density of prey. The number of prey consumed daily by each predator stage tested, increased more steeply at lower than at higher prey densities, exhibiting thus the type 2 functional response

---

Résumé Consommation des larves et des ♀ d'Aphis craccivora par les11^e stades, les 4^e stades et les adultes deCheilomenes vicina a été étudiée à une température variant de 24 à 30°C. Les jeunes stades du puceron sont consommés en nombre plus grand significativement que les derniers stades et les ♂. Les taux d'alimentation deC. vicina manifestent une corrélation hautement significative avec la densité de population de la proie. Le nombre de proies consommées quotidiennement par chaque stade considéré du prédateur augmentait plus brusquement aux faibles qu'aux fortes densités de proies, manifestant ainsi chez le prédateur le type 2 de réponse fonctionnelle.

     

Responses of Schizaphis graminum (Homoptera: Aphididae) to leaf excision in resistant and susceptible sorghum

Greenbugs, Schizaphis graminum (Rondani), were reared on intact and excised leaves of varieties of sorghum which differed in their suitability as hosts for this aphid. Aphids grew poorly on intact leaves of three resistant varieties, but grew well on excised leaves of the same varieties. Leaf excision did not affect aphid growth on three susceptible varieties. By electronically monitoring the feeding behaviour of aphids on two resistant and one susceptible variety, significant differences were found in many parameters between aphids assayed on excised vs. intact leaves of only the resistant varieties. Aphids on excised leaves of the resistant varieties, and on excised or intact leaves of the susceptible variety, made fewer probes to the phloem, spending more time ingesting from phloem during each probe, compared to aphids on intact resistant plants. There was a higher level of free amino acids in excised leaves of all varieties, but aphid growth and feeding behaviour improved as a result of excision only on resistant varieties. This observation, coupled with the fact that intact plants of all varieties have similar amino acid levels, indicates that these nutrients are not of primary importance in sorghum suitability to the greenbug. Other explanations for the aphids' responses to excised leaves are discussed.     

Effect of Manipulations on the Host Selection Behavior of Sitobion avenae (Homoptera: Aphididae)

The study of aphid host selection and feeding behavior is difficult because aphids have to penetrate the plant to reach their feeding site, phloem tissue. The activity of the stylets, salivation or food intake, can not be observed externally and requires an indirect visualization technique such as the Electric Penetration Graph (EPG). The plant selection behavior of Sitobion avenae on potato varied depending on whether an ethological or EPG method was used to study it. A similar variation did not occur with Myzus persicae or Rhopalosiphum padi. The application of water-based silver conductive paint onto the thorax, as normally used for EPG, or onto the abdomen of Sitobion avenae alates resulted in increased duration and frequency of probing compared to results from ethological observations. Our results indicated that EPG manipulations might have different effects on different species of aphids and that a comparison of EPG and ethological data is required to confirm that the EPG method does not bias aphid feeding behavior.     

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.     

Demographic parameters of cotton aphid, Aphis gossypii Glover （Homoptera： Aphididae） on five cotton cultivars

Life table parameters of the cotton aphid, Aphis gossypii Glover, were estimated on five cotton cultivars （‘ Sealand＇ ,‘Siokra＇ ,＇ Vararnin＇ ,‘ Bakhtegan＇ and ‘ Sahel＇ ）. Demographic parameters of the cotton aphid were assessed at 27.5 ± 1 ℃, 65% ＋ 10% RH and a photoperiod of 14：10 （L： D）h. The shortest developmental time for the nymphal stages was 5.5 days on ‘Siokra＇ and the longest was 6.1 days on‘ Sealand＇. The highest offspring per female was 29.6 on ‘Sahel＇ and the lowest was 15.3 on ‘Sealand＇. The rm values varied from 0.272 on ‘Sealand＇ to 0.382 （day^-1） on ‘Varamin＇. Jackknife estimates of the A. gossypii parameters on these cultivars indicated the greatest developmental rate and fecundity on ‘ Varamin＇ and the poorest on ‘ Sealand＇ cultivar.     

Effects of treatments for electrical penetration graph recordings on behaviour and biology of Aphis craccivora (Aphididae)

Abstract. Two laboratory studies were conducted to investigate effects of treatments for direct "current electrical penetration graph (DC-EPG) recordings or 'tether effect', on behaviour and reproductive performance of cowpea aphid Aphis craccivora Koch. The experiments constituted a control study in application of DC-EPGs to analyse cowpea aphid feeding behaviour and host plant resistance mechanisms. Resistant (ICV-12) and susceptible (ICV-1) cultivars of cowpea Vigna unguiculata (L.) Walp were used. EPG treatments included two groups of aphids: tethered aphids that were exposed to DC electricity via an attachment of a thin, flexible gold wire on their dorsum using a droplet of adhesive silver paint, and 'free' (untethered) aphids with a dorsal spot of silver paint only. EPGs of the tethered aphids were recorded continuously for c. ! h, whereas from the 'free' aphids recordings were done only for brief periods of 2–5 min, by temporarily contacting a gold wire to the spot of silver paint. Waveform signals generated from resistance fluctuations and electromotive forces, and representing aphid stylet penetration behaviour were recorded. A separate experiment was conducted to investigate effects of EPG treatments on aphid survivorship and population growth. Overall, EPG treatments did not significantly affect aphid stylet penetration behaviour or life-table parameters. However, effects of crop cultivar on those characteristics were significant. Waveform E2, which denotes aphid ingestion in phloem sieve elements, and non-penetration behaviour were important indicators of aphid resistance in ICV-12. Also, apart from the number of aphid generations, other life-table parameters were useful indicators of ICV-12 resistance. Thus, DC-EPGs provided a reliable technique for studying aphid stylet behaviour, and investigation of aphid resistance in cowpeas.     

Seasonal dynamics of cereal aphids on Russian wheat aphid (Homoptera: Aphididae) susceptible and resistant wheats

Field experiments were conducted in 1997 and 1998 to evaluate the impact of resistance to Russian wheat aphid, Diuraphis noxia (Mordvilko), on the cereal aphid complex in wheat. Two spring wheats were planted: the variety "Centennial" (Russian wheat aphid susceptible) and the advanced line IDO488 (Russian wheat aphid resistant). IDO488 incorporates the resistance found in PI 294994 into a Centennial background. Field plots were artificially infested with adult D. noxia and sampled weekly. The most abundant aphid species in 1997 were Metopolophium dirhodum (Walker), Sitobion avenae (F.), D. noxia, and Rhopalosiphum padi (L.). In 1998, the order of abundance was M. dirhodum, R. padi, S. avenae, and D. noxia. The resistant genotype had significantly fewer D. noxia than the susceptible one during both years. However, plant genotype had no significant effect on the other aphid species in either year. Both the initial density of D. noxia and plant growth stage, had a significant effect on D. noxia population development, but had no effect on the other aphid species. There was no interaction between D. noxia resistance and the population density of the other aphid species observed.     

Functional response of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) to Aphis gossypii Glover (Homoptera: Aphididae) in the Laboratory

Stage specific functional response of Harmonia axyridis (Pallas) to varying densities of Aphis gossypii Glover was examined in a simplified cucumber leaf arena under laboratory conditions. All stages of H. axyridis were isolated individually for 24 h with different prey densities at 25 °C and a photoperiod of 16:8 (L:D) h. The number of prey consumed by the predator was checked at 3, 6, 12, and 24 h. All stages of H. axyridis showed a Type II functional response. Based on the random predator equation, estimated attack rates of H. axyridis at 24 h were 0.0037, 0.0442, 0.3590, 0.3228, and 0.1456, and estimated handling times were 4.1001, 2.4575, 0.7500, 0.2132, and 0.1708 h for the first, second, third, and fourth instars, and female adult, respectively.     

Electrical penetration graphs of the damson-hop aphid, Phorodon humuli on resistant and susceptible hops (Humulus lupulus)

Electrical penetration graphs (EPG's) were used to locate resistance to Phorodon humuli (Schrank) (Homoptera, Aphididae) in hops (Humulus lupulus, Cannabinaceae). Aphids on those hops showing resistance had a much reduced E2 pattern (uptake of phloem). In addition, many aphids on the resistant plants spent time non-probing within two minutes of withdrawing from the phloem. This was not observed with aphids on susceptible hops. The results suggest that resistance is located in the phloem. The involvement of a mechanical factor such as the blocking of aphid stylets, the presence of antinutritional factors, or simply an inadequate supply of nutrients, are discussed as possible resistance mechanisms.     

Impact of density of Aphis craccivora (Aphididae) on growth and yield of susceptible and resistant cowpea cultivars

Impact of initial density of cowpea aphid Aphis craccivora Koch (Aphididae) at infestation on the growth and yield of aphid-susceptible cowpea cultivar ICV-1 and aphid-resistant cultivar ICV-12, was investigated. Plants at the seedling, flowering and podding stages of development were infested with five aphid densities consisting of 0, 2, 5, 10 and. 20 aphids per plant and maintained for 22 days. Extended leaf heights of plants and aphid counts were recorded at 7, 12, 17 and 22 days after infestation. Two crop growth parameters (biomass duration and leaf area duration), and two plant yield parameters (number of pods per plant and number of seeds per pod) were recorded. Due to the occurrence of parthenogenesis and changes in population dynamics during infestations, aphid densities were converted into cumulative cowpea aphid-days, to facilitate data analyses and interpretation. ANOVA indicated that there was significant (P=s 0.05) difference in aphid-day accumulations between the two cultivars when infested at the seedling stage. Accumulations on cv. ICV-1 were greater than on cv. ICV-12. However, no such differences between the cultivars were detected when plants were infested at flowering and podding stages. Therefore, the seedling stage was used for comparisons of the impact of cowpea aphid-days on the growth and yield parameters of the two cultivars. At the 95% confidence intervals, ICV-12 plants were consistently taller than ICV-1 plants. Infested ICV-1 seedlings showed stunting and other growth deformities which were not observed on ICV-12 plants. Regression analyses revealed substantial reductions in the growth and yield parameters of ICV-1 relative to ICV-12. Overall, cowpea aphid-days provided a convenient and reliable method for studying the aphid population dynamics and the subsequent impact on plant growth and yield performance.     

Suitability of Aphis fabae, Myzus persicae (Homoptera: Aphididae) and Aleyrodes proletella (Homoptera: Aleyrodidae) as prey for Coccinella undecimpunctata (Coleoptera: Coccinellidae)

The suitability of Aphis fabae Scopoli, Myzus persicae Sulzer and Aleyrodes proletella L. as food sources for Coccinella undecimpunctata L. was evaluated by studying the impact of prey consumption on the predator’s population growth parameters and feeding parameters. Unlike A. proletella, A. fabae and M. persicae supported the development and reproduction of C. undecimpunctata. A. fabae and M. persicae were considered to be essential prey, whereas A. proletella was considered to be an alternative prey. Aphid species showed different degrees of suitability: M. persicae significantly decreased the pre-oviposition period and increased adult longevity, fecundity and fertility compared with A. fabae. Moreover, A. fabae represents a suitable diet for larval development, but is not a suitable food source for adult reproduction. The predator’s population growth parameters, R₀, r_m and λ were increased with M. persicae, whereas T decreased. We found that the 4th instar larvae were the most voracious, particularly when fed on M. persicae; nevertheless, with this prey daily weight gain and feeding efficiency of 4th instar larvae were similar to that of individuals fed with A. fabae.     

Infection of Sentinel Cotton Aphids (Homoptera: Aphididae) by Aerial Conidia ofNeozygites fresenii(Entomophthorales: Neozygitaceae)

Uninfected adultAphis gossypii(Homoptera: Aphididae) apterae (sentinel aphids) on cotton leaves were exposed for 8 h to the air over a commercial cotton field in Louisiana during the night of 1–2 July 1995. At 0015 h there were 90,437 primary conidia/m³air of the fungusNeozygites fresenii(Entomophthorales: Neozygitaceae) at the midfield position as determined from Rotorod samples. Forty-eight percent (n = 106) of the sentinel aphids exposed for 8 h at midfield were infected by aerial conidia ofN. fresenii.Exposure of sentinel aphids outside the cotton field, at 10 and 100 m downwind and 10 m upwind, resulted in 34.8% (n = 131), 24.0% (n = 129), and 17.4% (n = 146) infected aphids, respectively. These data demonstrate that wind-dispersed aerial conidia ofN. freseniiare infective and rapidly and efficiently disperse the pathogen throughout aphid populations within and between fields.     

Parasitoid-host relationship betweenTrioxys (Binodoxys) Indicus [Hymenoptera: Aphidiidae] andAphis Craccivora [Hemiptera: Aphididae]. VI. Impact of males on the number of progeny of the parasitoid reared on certain host plants

The present paper elucidates the impact of males on the reproductive rate ofTrioxys (Binodoxys) indicus reared on the aphids ofCajanus cajan, Dolichos lablab andSolanum melongena host plants. The F₁ offspring increases with the increase of parasitoid number. This increase is maximum inC. cajan followed byD. lablab andS. melongena reared aphids. The presence of male parasitoids caused a significant decrease in the emergence of F₁ generation.      

Reproductive compatibility and genetic variation between two strains of Aphelinus albipodus (Hymenoptera: Aphelinidae), a parasitoid of the soybean aphid, Aphis glycines (Homoptera: Aphididae)

Aphelinus albipodus Hayat and Fatima is a potential biological control agent of the soybean aphid, Aphis glycines Matsumura, which is a newly introduced soybean pest in the United States. We compared the reproductive compatibility and molecular genetic variation between two geographic strains of A. albipodus. One strain was collected from soybean aphids in Japan and the other recovered from Russian wheat aphid, Diuraphis noxia (Mordvilko), in the western U.S., populations of which were established with parasitoids imported from Eurasia. We present results of crossing experiments between the two strains, genetic differences based on RAPD-PCR markers, rDNA ITS1 and ITS2 gene sequences, and presence of Wolbachia in the two strains using PCR amplification of the wsp gene. We found no reduction in the production of females in reciprocal crosses between strains, but a significant reduction in fecundity when F₁ females stemming from one of the reciprocal crosses were backcrossed to males from either source. The two strains differed by 3.4% in the rDNA ITS1 sequence and by presence/absence of one RAPD-PCR marker from a total of 20 RAPD primers screened, but their rDNA ITS2 sequences were identical. We used restriction enzyme analysis to separate the strains by differential digestion of the ITS1 PCR product. Wolbachia was present in 100% of males and females of both strains of A. albipodus.     

Classical biological control of Aphis gossypii (Homoptera: Aphididae) with Neozygites fresenii (Entomophthorales: Neozygitaceae) in California cotton

In August 1994 and 1995 classical biological control releases were made in cotton in the San Joaquin Valley, California, with an Arkansas strain of the entomopathogenic fungus, Neozygites fresenii, a pathogen of the cotton aphid, Aphis gossypii. Pre-release samples in both years indicated that N. fresenii was not naturally present in A. gossypii populations in the San Joaquin Valley. Two release methods were compared: dried N. fresenii-infected cotton aphid “cadavers” and chamber inoculation of A. gossypii. Both methods were successful in introducing N. fresenii to cotton aphids in California; however, higher prevalence of fungal infection resulted with the cadaver treatments. N. fresenii persisted and spread in the aphid population until early October 1994 and late September 1995. The highest mean percentage infection in the cadaver treatment in 1994 reached a level (14%) considered imminent for epizootics (12–15%). The use of predator exclusion cages resulted in higher N. fresenii prevalences.     

不同类型品种棉花上棉蚜适生性及种群动态

通过田间调查、室内网罩盆栽苗测定选择性等方法,考察了常规棉（泗棉3号,石远321）、杂交抗虫棉（辽棉19号,鲁棉研18号）、转单价基因抗虫棉（国抗12号,中棉所32）和转双价抗虫棉（SGK321,中棉所41）4种类型8个品种棉花上棉蚜的适生性及种群动态。结果表明： 棉蚜在各棉花品种上的种群动态有明显差异（P<0.05）,单株蚜量以转单价基因抗虫棉中棉32上最高,常规棉泗棉3号上最低,分别为297.81头/株和76.88头/株。棉蚜对4种类型棉花品种的选择性有明显差异（P<0.05）,其中对转单价基因抗虫棉有很强的选择性。根据棉蚜实验种群的参数判断,其在不同品种棉花上的生长发育、存活及繁殖存在显著差异： 若虫发育历期常规棉石远321上最长（6.46天）,双价棉中棉所41上最短（5.75天）; 存活率转单价基因抗虫棉中棉32上最高（88.21%）,双价棉SGK321上最低（76.46%）; 单雌产蚜量杂交抗虫棉辽棉19上最大（44.48头）,双价棉SGK321上最小（33.51头）; 内禀增长率转单价基因抗虫棉中棉32上最高（0.3695）,双价棉中棉所41上最低（0.3389）。综合评价,棉蚜的生存和繁殖适合性在转单价基因抗虫棉上最高,在双价棉上最低。     

基于EPG的麦长管蚜、麦二叉蚜和禾谷缢管蚜取食行为比较

麦长管蚜Sitobion avenae (Fabricius),麦二叉蚜Schizaphis graminum (Rondani)和禾谷缢管蚜Rhopalosiphum padi (Linnaeus)虽然都取食小麦,但其取食部位、传毒能力均有差异。利用EPG研究了3种麦蚜在同一寄主植物上的取食行为,结果显示:(1)禾谷缢管蚜在取食过程中G波出现的时间最早,G波总的持续时间和每次刺探G波的平均持续时间都明显长于麦长管蚜和麦二叉蚜。麦长管蚜第1次E1波的时间、E1波总的持续时间和每次刺探E1波的平均持续时间都明显比麦二叉蚜和禾谷缢管蚜要长。麦二叉蚜E2波总的持续时间和每次刺探E2波的平均持续时间都明显长于麦长管蚜和禾谷缢管蚜。表明3种蚜虫各自拥有不同取食策略。禾谷缢管蚜比其它两种蚜虫需要更多的水分和无机盐,麦长管蚜通过分泌大量唾液增大对食物的利用率,麦二叉蚜通过增加韧皮部取食时间以满足对营养的需要。(2)刺探过程中麦二叉蚜pd波出现的次数为(51.97±7.82)次,高于麦长管蚜的(44.73±4.52)次和禾谷缢管蚜的(32.99±4.22)次。麦二叉蚜pd波Ⅱ-2和Ⅱ-3亚波的时间最长,禾谷缢管蚜最短,三者之间差异达显著水平。表明与其它两种麦蚜相比,麦二叉蚜在取食过程中口针与细胞间的接触频率更高,与细胞内获毒和传毒相关的Ⅱ-3波和Ⅱ-2波持续时间更长,增加了获毒和传毒的几率,是其传毒能力最强的机理。