AGGREGATION IN APHIS FABAE SCOP. I. AGGREGATION ON PLANTS

A study of free infestations of Aphis fabae Scop. on the leaves of its host plants, spindle and sugar beet, showed that the clustered distribution of the aphids within the boundaries of each leaf was not a matter of chance but involved active aggregation. Experiments in small leaf cages showed that aggregation could take place independently of any differences between portions of a leaf, being brought about by truly gregarious reactions between the aphids. Experiments in which walking aphids were made to encounter other live or dummy aphids fixed to leaves, showed that the gregarious reactions were of at least two kinds. The aphids both probed more often, and remained inserted for longer, near other aphids than apart from them.
It appears that the unequal distribution of aphids among the leaves of a plant is due primarily to intrinsic differences between leaves, but is aided by gregariousness, while the aggregations observed on leaves are due primarily to gregariousness, and only secondarily to differences between portions of a leaf. It is suggested that the primary function of gregariousness is to concentrate the aphids on to the best feeding sites, both among the parts of a plant, and among whole plants; that is, it acts as an aid in host selection, even with aphids which do not aggregate so compactly as A. fabae. The functions of compact aggregation and other specialized forms of gregarious behaviour remain to be discovered.     

Effect of honeydew and hosts on plant colonization by the aphid parasitoidEphedrus cerasicola

Females ofEphedrus cerasicola Stary were released into small glasshouses (13.6 m³) or cages (0.125 m³) which contained paprika plants in 2 of the following categories: fresh plants (without honeydew and aphids), honeydew-contaminated plants (aphids removed) and aphid-infested plants (with honeydew). Two hundred females were released into each glasshouse with 20 plants, 10 of each category, while 10 females were released into the cages with 2 plants, one of each category. Control experiments with only fresh plants were conducted in both the glasshouses and the cages. The female parasitoids on each plant were counted at regular intervals for 24 or 48 h in the glasshouse and for 4 h in the cage experiments; this number was used as an indicator of plant preferences. The following preferences were established (preferred plant > less preferred plant): honeydew-plant > fresh plant, aphid-plant > fresh plant, aphid-plant > honeydew-plant. The number of females on the aphid-plants increased throughout the experimental period, whereas the number on the honeydew-plants tended to be more stable.      

A simple,light clip‐cage for experiments with aphids

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Effects of prey density and spatial distribution on prey consumption of the adult predatory ladybird beetle

The effects of prey density and spatial distribution on prey consumption of the adult predatory ladybird, Harmonia axyridis , were investigated by using a 2 × 2 factorial design in large scale cages. Prey density influenced prey consumption of the ladybirds, and the frequency with which predation occurred was quite different between the prey distributions. The ladybirds consumed a relatively constant and small number of aphids when the prey were uniformly distributed, whereas the number of prey consumed per day when predation occurred was large and much more variable when the prey were contagiously distributed. At high prey density, the number of prey consumed was highest during the first day of the experiment; thereafter, only 10–20 aphids were consumed during the following 3 days. However, these patterns of prey consumption were not observed at low prey density. The percentage of aphids that remained on the host plants when the experiments were terminated was higher at low prey density than at high prey density, suggesting that predator foraging efficiency at low prey density was lower than at high prey density. Ladybirds foraging for high prey density were more frequently observed on the plants with aphids than ladybirds foraging for low prey density. Prey distribution also influenced the frequency of residence of ladybirds on the plants. The different predation patterns observed in the two spatial distributions, in which prey consumption was much more variable for the contagious distribution, might be explained by the difference in prey encounter rate of the predator between the distributions. This study indicated that the ladybirds had limited ability to search out prey over large spatial scales.     

3种寄主上桃蚜的选择性及形态分化

桃蚜是一种重要的农业害虫,寄主广泛,种下分化复杂。以采自黄土高原旱作区桃树、烟草、甘蓝上的桃蚜为研究对象,通过叶片选择法、传统比较形态测定法研究了3种寄主上桃蚜的选择性及形态分化。结果表明:在3种寄主同时存在的情况下,烟草上的桃蚜嗜食烟草,表现为63.5%的桃蚜选择烟草叶,13.8%选择甘蓝叶,8.2%选择桃叶,而甘蓝和桃树上的桃蚜对原寄主没有表现出明显的嗜好性;从形态指标来看,3种寄主上的桃蚜在体长、触角末节长度、后足腿节长度、触角与体长的比例方面存在显著差异(P0.05),说明这几个特征可以作为区分这3种寄主上桃蚜的依据。综合分析可以初步认为黄土高原旱作区烟草上的桃蚜可能形成了寄主专化型-烟草型。     

Methods of Recording Aphid Populations for Use in Research on Potato Virus Diseases

Methods of counting aphids infesting potato crops are reviewed and discussed. The results of an aphid survey in different parts of England during 1946 and of counts at Rothamsted during 1947 are given and used to illustrate methods of expressing the counts; the extent of virus spread in 1946 showed no close correlation with aphid numbers. It is concluded that an estimate of the number of aphids per plant is better for virus disease research than the number per 100 leaves. A method of estimating the number of aphids per plant with tolerable accuracy is described, but it is concluded that for most surveys a method which does not necessitate the counting of the aphids is adequate. Such a method, based on the percentage of leaves infested, is described.     

Increased reproduction by pea aphids in the presence of secondary parasitoids

1. The influence of the presence of secondary parasitoids on aphid reproduction was tested in a field experiment. 2. Single pea aphids Acyrthosiphon pisum in clip cages on potted bean plants were exposed to a collection of secondary parasitoids enclosed within a small perforated bag. The aphids were exposed to volatile chemicals released by the secondary parasitoids but there was no physical contact. 3. The production of nymphs was recorded over an 11‐day period. Aphids produced significantly more offspring in the presence of secondary parasitoids than in the absence of secondary parasitoids (36.9 vs. 31.2). 4. This is the second reported occurrence of this phenomenon, and the reasons for its occurrence and its consequences for aphid population and community ecology are discussed.     

Soybean cyst nematode effects on soybean aphid preference and performance in the laboratory

Herbivores on plants frequently interact via shared resources. Studies that have examined performance of herbivores in the presence of other herbivores, however, have often focused on above-ground feeding guilds and relatively less research has examined interactions between below- and above-ground consumers. We examine how soybean aphid, Aphis glycines (Matsumura) an above-ground phloem-feeding herbivore, interacts with a below-ground plant parasite, soybean cyst nematode, Heterodera glycines (Ichinohe) through their shared host plant, soybean (Glycine max L). Laboratory experiments evaluated the preference of alate (flight-capable) soybean aphids toward plants either infected with soybean cyst nematode or uninfected controls in a simple choice arena. Alate soybean aphids preferred uninfected soybean over soybean cyst nematode-infected plants: 48 h after the releases of alate aphids in the center of the arena, 67% more aphids were found on control soybean compared with nematode infected plants. No-choice feeding assays were also conducted using clip cages and apterous (flight-incapable) aphids to investigate effect of soybean cyst nematode infection of soybean on aphid performance. These studies had mixed results: in one set of experiments overall aphid population growth at 7 d was not statistically different between control and soybean cyst nematode-infected plants. A different experiment using a life-table analysis found that apterous aphids feeding on soybean cyst nematode-infected plants had significantly greater finite rate of increase (λ), intrinsic rate of increase (r(m)), and net reproductive rate (R(o)) compared with aphids reared on uninfected (control) soybean plants. We conclude that the below-ground herbivore, soybean cyst nematode, primarily influences soybean aphid behavior rather than performance.     

Scale-dependent indirect interactions between two prey species through a shared predator

We investigated the potential for indirect interactions between two prey species, pea aphids ( Acyrthosiphon pisum ) and potato leafhoppers ( Empoasca fabae ), through a shared predator (Nabis spp.), and how these interactions may change across three spatial scales. In greenhouse experiments using small clusters of plants containing pea aphids and/or potato leafhoppers, the predation rates on both pea aphids and potato leafhoppers were independent of the presence of the other species, indicating no indirect interactions. In greenhouse experiments using cages containing 48 plants, when aphids and leafhoppers were confined to separate plants among which nabids could move, pea aphids had a positive effect on the survival of potato leafhoppers from predation. The positive effect of aphids on leafhoppers occurred because nabids spent more time on plants harboring aphids, thereby drawing nabids away from plants containing leafhoppers. Finally, we measured the abundance of nabids in a large-scale experiment designed to manipulate the abundances of pea aphids and potato leafhoppers in alfalfa fields. Fields with high aphid density contained more nabids, thereby suggesting that pea aphids will have a negative indirect effect on potato leafhoppers by increasing the density of nabids within fields. Potato leafhoppers had no indirect effects on pea aphids at any scale. This study shows that indirect interactions between prey species may depend upon spatial scale, because the factors affecting a predator's diet choice on a small scale may differ from those factors affecting a predator's distribution at larger scales.     

Pairwise interactions between functional groups improve biological control

Ecologists have long debated whether predators primarily disrupt one another’s prey capture through interspecific interference, or instead complement one another by occupying different feeding niches. Resolution of this debate has been difficult because different experimental designs are typically used to study interference versus complementarity. We adopted a somewhat atypical approach, surveying communities of predatory insects on 73 free-growing Brassica oleracea plants, and then re-constructing each community in field cages to measure its impact on aphid prey. The predator communities naturally varied in species composition, richness, and relative abundance; in our experiment we kept total predator density constant to avoid confounding effects of differing overall abundance. The predator communities’ impacts on aphids differed by >10-fold. Using a generalized linear model, we found that pairings of several predators in the community improved aphid suppression while no pairings disrupted it. Indeed, accounting for the presence of the beneficial pairings provided more power than species richness to explain predators’ impacts on aphids. Altogether, our results suggest generally complementary or neutral, rather than disruptive, multi-predator effects in this community. Our approach may be useful for determining the frequency of complementary species-pairings in many other systems.     

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.     

The impact of Meu1-mediated resistance in tomato on longevity, fecundity and behavior of the potato aphid, Macrosiphum euphorbiae

The effect of the tomato resistance gene, Meu1, on feeding, longevity, fecundity and developmental rate of the pink biotype of the potato aphid, Macrosiphum euphorbiae (Thomas) (Hemiptera, Aphididae), was determined using nearly isogenic tomato (Lycopersicon esculentum Mill, Solanaceae) lines. Aphid mortality was significantly higher on resistant plants, with 60% of the aphids dying by the 4th day of exposure. By the 10th day, all the aphids on the resistant plants were dead whereas 100% of the aphids on susceptible plants were alive. Meu1-mediated resistance resulted in significantly decreased fecundity with a ten-fold decrease in the net fertility rate (4.5 and 45.7 progeny per aphid on resistant and susceptible tomato, respectively). A qualitative analysis showed that honeydew was produced by aphids on resistant and susceptible plants, suggesting that aphids initiate feeding on both plant types. However, significantly lower quantities of honeydew were present when aphids were caged on resistant plants. There were also significant differences in aphid location on resistant and susceptible leaves. Experiments evaluating behavior in less than 24 h showed that aphids left resistant leaves after relatively short exposure (3–6 h). Aphids transferred from resistant to susceptible tomato at intervals between 3 h and 24 h resumed feeding as evidenced by presence of honeydew. Although the mechanism by which Meu1-mediated resistance operates is not yet known, our data suggest that resistance factors act rapidly after initiation of feeding and that lower fecundity and longevity are related to reduction in aphid feeding.     

THE DIFFERENT DISTRIBUTION OF TWO BRASSICA VIRUSES IN THE PLANT AND ITS INFLUENCE ON SPREAD IN THE FIELD

Previous knowledge provided no explanation for the greater prevalence of cauliflower mosaic than of cabbage black ring spot in field crops of cauliflower. Both viruses are spread principally by Myzus persicae and Brevicoryne brassicae , and both are transmitted equally readily from infected seedlings. Cabbage black ring spot virus has a much wider host range, and sap from infected leaves has a higher dilution end-point than sap from leaves infected with cauliflower mosaic virus.
At least part of the difference between the rate at which the two viruses spread in the field may be accounted for by the different manner in which they are distributed in old infected plants, and the effect this has on transmission by aphids. Cauliflower mosaic virus occurs in high concentration in all the new leaves produced by infected plants. Cabbage black ring spot virus, on the other hand, occurs mainly in the older leaves, and even there is localized in parts that show symptoms. Only in recently infected plants does cabbage black ring spot virus occur in young leaves.
After flying, most aphids alight on the upper parts of plants; they are therefore less likely to acquire cabbage black ring spot virus than cauliflower mosaic virus. It may be significant that cabbage, a host in which old leaves are in a more favourable position for alighting aphids than are those of cauliflower, is also often extensively infected with cabbage black ring spot virus.     

Location of resistance factors in the leaves of potato and wild tuber-bearing Solanum species to the aphid Myzus persicae

Analysis of electrically recorded feeding behaviour of aphids was combined with colony‐development tests to search for sources of resistance to Myzus persicae (Sulzer) (Homoptera: Aphididae) in tuber‐bearing Solanum species (Solanaceae), aiming at a reduction of potato leaf roll virus (PLRV) transmission. Twenty genotypes, originating from 14 gene bank accessions, representing 13 wild tuber‐bearing Solanum spp., three Solanum tuberosum L. (potato) cultivars, and one S. tuberosum breeding line, were selected. Colony‐development tests were carried out in no‐choice experiments by placing adult aphids on plants of each genotype and counting numbers of nymphs and adults on young plants after 8 and 15 days, and on flowering plants after 14 and 30 days. Large differences were observed among genotypes: some developed small colonies and others developed large ones. Also, in a few genotypes, resistance in mature plants was different for leaves of different ages; young leaves were resistant to aphids whereas old senescent leaves were susceptible. The electrical penetration graph (DC‐EPG system) technique was used to study aphid feeding behaviour on each Solanum genotype for 6 h. Electrical penetration graph (EPG) results also showed large differences among the genotypes, indicating resistance at the leaf surface and at three different levels of plant tissue (epidermis, mesophyll, and phloem). Therefore, it was concluded that different mechanisms of resistance to M. persicae exist among the genotypes analysed. EPGs recorded from aphids on Solanum berthaultii Hawkes and Solanum tarijense Hawkes with and without glandular trichomes showed that strong surface resistance can bias EPG parameters associated with resistance located in deeper tissues. Experimental evidence is presented that the resistance to aphids in the genotypes with glandular trichomes strongly depends on these morphological structures.     

Changes in photosynthetic and transpiration rates of cotton leaves infested with the cotton aphid, Aphis gossypii: Unrestricted infestation

The influence of aphid, Aphis gossypii, feeding on photosynthesis and transpiration in cotton plants was investigated under greenhouse conditions. Four population densities of 0, 5, 10, and 25 aphids were used to infest individual cotton leaves. Gas exchange rates were determined for single attached cotton leaves after 9, 18, and 27 days of aphid infestation. Aphid feeding changed photosynthetic rates and transpiration rates. These changes were proportional to the number of aphids and the length of infestation period. Photosynthetic rates were significantly reduced in infested leaves with 25 aphids over 18 days, whereas significant reduction in photosynthetic rates was recorded within 27 days in infested leaves with 5, 10, and 25 aphids in comparison to their respective control. Initial population of 10 aphids increased significantly the transpiration rate of infested leaves over 9 and 27 days. Leaves of plants with 25 aphids had significantly greater transpiration rate than the control at all times.     

The effect of cereal growth stages on the reproductive activity of Sitobion avenue and Metopolophium dirhodum

Apterous Sitobion avenae on oats were found to have a higher reproductive rate on the ears (5–95 nymphs per day) than on young leaves (3–78 nymphs per day), mature leaves (2–17 nymphs per day) or senescent leaves (2–08 nymphs per day). At each growth stage of the the host plant large aphids gave birth to more young than small aphids, but aphids of the same weight on different growth stages of the host plant had different reproductive rates. On wheat in the field S. avenae gave birth to more young on the ears but less on the leaves than Metopolophium dirhodum on the leaves.
The suitability of ears to S. avenae changed through the flowering and ripening stages. During the latter part of the milky-ripe stage the reproductive rate of S. avenae declined and its mortality increased. The consequences of the variations in reproduction and mortality are discussed in relation to the life history and population dynamics of S. avenae and the assessment of cereal varieties for aphid resistance.     

烟粉虱(Bemisia tabaci)的寄主选择性

通过田间系统调查、实验室嗅觉测定、笼内和培养皿内自由扩散观察,对烟粉虱的寄主选择性进行了研究.结果表明,在田间,烟粉虱对不同的寄主植物存在明显的寄主选择性,其中对茄子、花椰菜、黄瓜等植物具有较强的嗜性,而对蕹菜、芹菜、苋菜等植物的嗜性较差;烟粉虱对同一植物的不同品种也有明显的选择性.烟粉虱对寄主植物颜色有明显的选择性,选择结果与烟粉虱对这些寄主的嗜性趋势基本一致.烟粉虱对单株寄主植物的嗅觉反应不敏感,但对植物叶片的乙醇抽提物有明显的嗅觉反应,并表现出较强的寄主选择性.在养虫笼内,烟粉虱从虫源皿向寄主植物自由扩散的过程中,在不同寄主植物和同一植物的不同品种上着落的成虫数量不同,并且在一定的时间范围内,着落在寄主叶片上的虫量还会不断的发生变化.     

The influence of aphids and honeydew on the leaving rate of searching aphid parasitoids from wheat plants

Dispersal cages were used to investigate the effects of aphids and treatment with artificial honeydew on the leaving rate of searching females of the parasitoid Aphidius rhopalosiphi from groups of wheat plants. Parasitoids which flew away from groups of plants placed in the centre of a cage were trapped on the sides and roof of the cage and thus were prevented from returning to the plants. The positions of trapped parasitoids suggested their direction of flight when dispersing from the plants. Parasitoids increased their residence times on groups of plants in the presence of aphids and of artificial honeydew, but the rate of parasitism of the host Sitobion avenae was not raised by the presence of artificial honeydew under the experimental conditions used. The direction of flight taken by the majority of parasitoids suggested that they were leaving the plants in order to locate further plants nearby to continue searching rather than to terminate searching and disperse away from the area. The need to consider plant patch size in studies of parasitoid searching behaviour is stressed.     

Quantitative studies on the uptake and retention of potato leafroll virus by aphids in laboratory and field conditions

Enzyme-linked immunosorbent assay (ELISA) was adapted for the efficient detection and assay of potato leafroll virus (PLRV) in aphids. Best results were obtained when aphids were extracted in 0.05 M phosphate buffer, pH 7.0, and the extracts incubated at 37 °C for 1 h before starting the assay. Using batches of 20 green peach aphids (Myzus persicae), about 0.01 ng PLRV/aphid could be detected. The virus could also be detected in single aphids allowed a 1-day acquisition access period on infected potato leaves. The PLRV content of aphids depended on the age of potato source-plants and the position of source leaves on them. It increased with increase in acquisition access period up to 7 days but differed considerably between individual aphids. A maximum of 7 ng PLRV/aphid was recorded but aphids more usually accumulated about 0.2 ng PLRV per day. When aphids were allowed acquisition access periods of 1–3 days, and then caged singly on Physalis floridana seedlings for 3 days, the PLRV content of each aphid, measured subsequently, was not strongly correlated with the infection of P. floridana. The concentration of PLRV in leaf extracts differed only slightly when potato plants were kept at 15, 20, 25 or 30 °C for 1 or 2 wk, but the virus content of aphids kept on leaves at the different temperatures decreased with increase of temperature. PLRV was transmitted readily to P. floridana at all temperatures, but by a slightly smaller proportion of aphids, and after a longer latent period, at 15 °C than at 30 °C. The PLRV content of M. persicae fed on infected potato leaves decreased with increasing time after transfer to turnip (immune to PLRV). The decrease occurred in two phases, the first rapid and the second very slow. In the first phase the decrease was faster, briefer and greater at 25 and 30 °C than at 15 and 20 °C. No evidence was obtained that PLRV multiplies in M. persicae. These results are compatible with a model in which much of the PLRV in aphids during the second phase is in the haemocoele, and transmission is mainly limited by the rate of passage of virus particles from haemolymph to saliva. The potato aphid, Macrosiphum euphorbiae, transmitted PLRV much less efficiently than M. persicae. Its inefficiency as a vector could not be ascribed to failure to acquire or retain PLRV, or to the degradation of virus particles in the aphid. Probably only few PLRV particles pass from the haemolymph to saliva in this species. The virus content of M. euphorbiae collected from PLRV-infected potato plants in the field increased from early June to early July, and then decreased. PLRV was detected both in spring migrants collected from the plants and in summer migrants caught in yellow water-traps. PLRV was also detected in M. persicae collected from infected plants in July and August, and in trapped summer migrants, but their PLRV content was less than that of M. euphorbiae, and in some instances was too small for unequivocal detection.