Movement by mating females of a host alternating aphid: a response to leaf fall

Autumn remigrants of the host alternating bird cherry-oat aphid, Rhopalosiphum padi , colonise individual plants of the winter host, Prunus padus . However, they do not select leaves at different stages of senescence, which is surprising as leaf fall is a definitive deadline for production and development of mating females. For successful development mating females should be mobile and have the capacity to evaluate leaf senescence. Although remigrants do not selectively colonise leaves at different stages of senescence, mating females were found in very low numbers on leaves that were about to be shed. In choice test and olfactometer bioassays, abscising leaves were avoided by mating females. In the field mature leaves were available for colonisation during the period of leaf fall of P. padus . We conclude that mating females abandon abscising leaves in favour of leaves at an earlier stage of senescence, increasing the time available for mating. Our results support the role of remigrants as pioneers which identify good winter host plants, and of mating females as mobile searchers for optimal feeding and mating sites.     

The life-cycle and host preferences of the bird cherry-oat aphid, Rhopalosiphum padi L., and their bearing on the theories of host alternation in aphids

Emigrants of the host-alternating bird cherry-oat aphid fly from the primary host to various grasses before the end of June, even in the absence of natural enemies on bird cherry. They fly before the leaves of bird cherry mature and at a time when numbers of insect predators are rapidly increasing. In the autumn gynoparae and males return to bird cherry whose senescing leaves again provide the aphids with a rich source of food. The growth efficiency of R. padi on bird cherry and oats indicates that the quality of the available food on the two hosts is markedly different.
Apterous exules prefer oats to young bird cherry leaves and survive best on oats. Gynoparae prefer bird cherry leaves and only successfully produce offspring on the primary host. This, and other work published on R. padi , support the premise that a change in the aphid's preferences causes the change of host.
The dual-discrimination theory of host alternation in aphids is examined in the light of this and other published work.     

The role of chemical cues in host finding and acceptance by Callosobruchus chinensis

We studied the response of female Callosobruchus chinensis to chemical cues emitted by cowpea seeds at different stages of bruchid infestation (uninfested, egg carrying, L1-, and L4-infested). Olfactory attractiveness was determined in Y-tube olfactometer assays by testing individual seed categories against either clean air or uninfested seeds. Oviposition preferences between uninfested and infested seeds were determined in petri-dish choice-experiments. The olfactometer assays revealed that weevils discriminate between seeds containing different stages of developing bruchids on the basis of olfactory cues. While odors from uninfested and egg-carrying seeds acted as attractants, odors from L1- and L4-infested seeds failed to induce a positive response by the bruchids. When given a choice between uninfested and infested seeds in the olfactometer, weevils preferred uninfested seeds over L1- and L4-infested seeds, but failed to distinguish between uninfested and egg-carrying seeds. In the oviposition experiment as well, bruchids showed a distinct preference for uninfested seeds when offered in combination with L1- and L4-infested seeds. This experiment further showed a reduced acceptance of egg carrying seeds. Our results indicate that C. chinensis females use chemical information during both host searching and host acceptance. Volatiles from uninfested or egg carrying seeds act as attractants, while deterrence increases as development of bruchid immature stages progresses.     

Response of the elm leaf beetle to host plants induced by oviposition and feeding: the infestation rate matters

We investigated by olfactometry and feeding‐ and oviposition‐choice‐tests how the highly specialised elm leaf beetle, Xanthogaleruca luteola Müller (Coleoptera: Chrysomelidae), responds to conspecifically induced defences in the field elm Ulmus minor Miller (Ulmaceae). While egg deposition of the beetle induced elms to release volatiles attractive to the egg parasitoid Oomyzus gallerucae Fonscolombe (Hymenoptera: Eulophidae), feeding alone did not. In the present study, females of the elm leaf beetle showed preferences for the odours of twigs induced by low egg deposition and feeding over odours from uninfested twigs. In contrast, heavy infestation rendered elm odours less attractive to the beetles. Feeding and oviposition bioassays revealed an oviposition preference for leaves from uninfested twigs when compared to locally infested leaves. However, beetles preferred to feed upon systemically induced leaves compared to uninfested ones. The different preferences of the elm leaf beetle during host plant approach might be explained by a strategy that accounts for both gaining access to high quality nutrition and avoiding competition or parasitism.     

Induced proteolysis within the bird cherry leaves evoked by Rhopalosiphum padi L. (Hemiptera, Aphidoidea)

The objectives of this study were to elucidate the impact of bird cherry-oat aphid (Rhopalosiphum padi L.) feeding on functioning of the proteolytic machinery in bird cherry leaves. Biochemical analyses proved that R. padi feeding in tissues of primary host stimulated activity of the two major fractions of proteinases (extracted at the optimal pH values: 5.0 and 7.0). Additionally, it has been demonstrated that aphids' feeding on bird cherry led to a decline in levels of albumins and globulins (main protein fractions in P. padus leaves). The opposite tendency, regarding the amounts of these protein fractions was ascertained at the phase of disappearance of R. padi population on tested shoots. Furthermore, it is reported that an increase in activity of the analysed enzymes and a decline in the content of tested protein fractions, were proportional to density of aphid individuals developing on P. padus side shoots. It is hypothesized that long-term R. padi feeding may lead to intensifying the catabolic processing of proteins by the activated proteolytic machinery in bird cherry leaves. The multi-level biological functions of endogenous plant proteinases and their significance in triggering the defense reactions in aphid-infested plant tissues are discussed.     

Nutritional enhancement of host plants by aphids - a comparison of three aphid species on grasses

Three aphid species were compared with respect to ability of enhancing the nutritional quality of their host plants. Rhopalosiphum padi, which does not induce macroscopic changes in its host plants, was compared with Schizaphis graminum and Diuraphis noxia, both of which induce distinctive types of chlorotic lesions. Phloem sap samples were collected from severed stylets of feeding aphids and from exudates of cut leaves of plants uninfested or infested with each aphid species. Samples were analyzed for concentrations of individual amino acids.Compared to R. padi, S. graminum ingested phloem sap with a two-fold higher concentration of amino acids and a much higher proportion of essential amino acids. Similar differences between these two aphid species were observed on both wheat and barley. For each aphid species, the absolute concentrations of amino acids and the relative proportions of essential amino acids were similar between the two host plants. Effects of D. noxia were similar to those of S. graminum, though less dramatic. Exudates from leaves infested with each aphid species showed relative concentrations of individual amino acids that were similar to those in the corresponding stylet exudates. Based on comparison of stylet exudates and cut leaf exudates from infested and uninfested plants, R. padi seems to have little effect on amino acid composition of phloem. Changes in the phloem induced by both S. graminum and D. noxia appear to be systemic, affecting at least the whole leaf they are feeding on. The changes observed for D. noxia and for S. graminum are likely to be nutritionally advantageous for the aphids and are expected to affect the aphids' dependence on nutritional supplementation by intracellular symbionts (Buchnera).     

Plant volatiles and density-dependent conspecific female odors are used by Asian citrus psyllid to evaluate host suitability on a spatial scale

We investigated how chemical cues derived from female Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae) and their host plants affect host acceptance choices by conspecifics. In four-choice cage and two-choice olfactometer assays, female psyllids avoided conspecific female cues in a density-dependent manner. However, odors from citrus plants actively damaged by psyllid feeding were attractive to conspecific females. When odors from feeding-damaged plants were presented simultaneously with odors from female D. citri, attraction of female conspecifics was no longer observed as compared with a clean air control in olfactometer assays. In subsequent experiments, D. citri females were released within arenas that contained actively feeding-damaged or non-damaged (control) citrus plants, each with previously psyllid-infested and uninfested young leaves. D. citri development is linked to the presence of these newly emerging leaves which is the only site of nymphal development. Female D. citri were initially attracted by the actively damaged plants as compared with non-damaged controls. After acceptance of plants that were actively damaged by feeding, D. citri females preferentially chose and settled on uninfested young leaves as compared with previously infested young leaves. A herbivore-induced plant volatile attractant and a female-specific odor repellent appear to be complementary foraging cues providing psyllids with information at two spatial scales: (1) the whole plant level for choosing a plant potentially harboring male conspecifics for mating, and (2) the within plant level to reduce intra-conspecific competition by identifying previously exploited resources.     

Change in response of Rhopalosiphum padi spring migrants to the repellent winter host component methyl salicylate

Olfactometry showed that the response of spring migrants of the bird cherry-oat aphid, Rhopalosiphum padi (L.) (Homoptera: Aphididae), to the repellent winter host volatile methyl salicylate changes with age of the adult aphid. Between three and four days after becoming adult, and having left the winter host Prunus padus L., aphids lost their negative response to the chemical. The change in response was not associated with contact with a summer host, oats. In a settling choice bioassay, migrants avoided oats which had been exposed to volatile methyl salicylate. Aphids with removed antennal tips did not avoid the exposed plant, indicating that plant choice was influenced by cues from the plant surface. The results are discussed in relation to the use of methyl salicylate in integrated control.     

The influence of the host plant of diamond-back moth (Plutella xylostella) on the plant preferences of its parasitoid Cotesia plufellae in Sri Lanka

Abstract. In Y-tube olfactometer tests, female Cotesia plutellae Kurdjmov (Hymenoptera: Braconidae), a major parasitoid of Plutella xylostella (L.) in southeast Asia, preferred the odour of the brassica type (Blue Lake cabbage) on which they had developed.Such plants were even preferred to the odour of caterpillar-infested and damaged leaves of another brassica (Early Phenomenal cauliflower).However, within one brassica type, the odour of caterpillar-infested leaves was preferred to that of uninfested and undamaged leaves.Odours of both Blue Lake and Early Phenomenal were preferred to the control odour (water).However, following the transfer of the parasitoid from Blue Lake to another cabbage (Brunswick) for one generation, females no longer distinguished between Blue Lake and Early Phenomenal.     

麦长管蚜和禾谷缢管蚜对小麦植株挥发物及蚜害诱导挥发物的行为反应

用四臂嗅觉计测定了麦长管蚜Macrosiphum avenae和禾谷缢管蚜Rhopalosiphum padi对小麦植株挥发物及麦蚜取食诱导挥发物的行为反应,揭示了2种麦蚜的嗅觉及小麦植株的诱导防御反应特点.在所选的13种小麦植株挥发物及蚜害诱导挥发物组分中,6-甲基-5-庚烯-2-酮、6-甲基-5-庚烯-2-醇和水杨酸甲酯对这2种蚜虫表现出强的驱拒作用;反-2-己烯醛对麦长管蚜的有翅和无翅蚜的吸引作用最强;反-2-己烯醇对禾谷缢管蚜的无翅蚜吸引作用最强,反-3-己酰醋酸酯对禾谷缢管蚜有翅蚜的吸引作用最强.说明麦蚜取食能诱导小麦植株的防御反应,麦长管蚜和禾谷缢管蚜及其不同蚜型间嗅觉反应的特点不同.     

Host choice by Aphidius colemani: effects of plants, plant–aphid combinations and the presence of intra-guild predators

Abstract.  1. This study first measured the effect of plant (radish, pepper, and wheat), aphid–plant combination (the green peach aphid Myzus persicae on pepper and radish, and the bird cherry-oat aphid Rhopalosiphum padi on wheat) and the host on which Aphidius colemani was reared (the cotton aphid Aphis gossypii on cucumber, M. persicae on radish and pepper, and R. padi on wheat) on host choice behaviour of the parasitoid and the performance of its offspring. Then, the effect of predator presence ( Coccinella undecimpunctata larvae) on host preference of the wasps was tested.
2. When reared on M. persicae on either radish or pepper , wasps preferred the aphid–plant combination from which they had emerged. Wasps reared on A. gossypii (naïve to all hosts offered) and R. padi preferred to parasitise M. persicae on radish and M. persicae on either radish or pepper, respectively. Rhopalosiphum padi on wheat was the least preferred and also the least suitable host, as determined by offspring body size.
3. Contrary to expectations, the presence of predators did not influence the host choice of A. colemani , even when predator-free plants were offered nearby. Results indicate that wasps are more likely to remain in some host systems but not in others, even when facing risk of predation.
4. Results are discussed in respect to field data and the relative importance of risk of predation and host preference to wasp fitness.     

The influence of the rearing host on the response of the parasitoid Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae) to odours from Ephestia kuehniella and Plodia interpunctella in a Y-tube olfactometer

Venturia canescens (Gravenhorst) is an ichneumonid generalist parasitoid that successfully attacks the larvae of different lepidopteran pests that infest stored products. These pest species include Plodia interpunctella and Ephestia kuehniella. In this study, we aimed to evaluate the influence of the rearing host on the parasitoid’s ability to detect and respond to a new host different from the rearing species. For this reason, the trials tested the preference of parasitoids reared on P. interpunctella or E. kuehniella for products that were or were not infested with larvae of these hosts. The trials were conducted in a Y-tube olfactometer. Regardless of the rearing host species, the parasitoids showed no preference for uninfested products. The parasitoids were attracted to products infested with larvae of their rearing host in preference to uninfested products. They also showed preferential attraction to products infested with the new host over uninfested products. E. kuehniella was the preferred host, irrespectively of the parasitoid host rearing species. The results are discussed to develop a better understanding of the ecology of V. canescens for its application in biological control.     

Olfactory response of the anthocorid predatory bug Orius sauteri to thrips-infested eggplants

The predator Orius sauteri (Poppius) (Heteroptera: Anthocoridae) is an effective arthropod natural enemy of thrips, especially Thrips palmi Karny (Thysanoptera: Thripidae), a serious pest of vegetables. First, we studied O. sauteri's response to the odour from thrips-infested eggplant [Solamum melongena L. (Solanaceae)] in a laboratory by two-choice experiments using a Y-tube olfactometer. When detached eggplant leaves were used as odour sources, O. sauteri preferred the volatiles from uninfested leaves to clean air. Concerning preferences among differently infested leaves, O. sauteri preferred the volatiles from plants infested with 10–100 thrips per leaf to uninfested leaves, but showed no significant preference for artificially damaged leaves over uninfested leaves. Similar results were obtained when complete plants were tested as the odour source. Second, release and recapture experiments in a greenhouse, a more realistic set of conditions, were conducted to confirm whether a significant preference for infested plants occurred at similar infestation levels as in the laboratory. Trends favouring infested plants were detected at densities of five and 500 thrips per plant; however, at the five thrips per plant, this trend was due to the large deviation seen in infested plants in only one replicate. In light of the low tolerable thrips density of eggplant, it is necessary to confirm whether artificial treatments with chemicals induce the emission of herbivore-induced plant volatiles that can attract O. sauteri and prolong its residence time on the leaf.     

Innate olfactory responses of female and male parasitoid Apanteles taragamae Viereck (Hymenoptera: Braconidae) toward host plant infested by the cucumber moth Diaphania indica Saunders (Lepidoptera: Crambidae)

The behavioural responses of Apanteles taragamae, a larval parasitoid of the cucumber moth Diaphania indica, to the volatiles of cucumber plants was investigated in a four-arm olfactometer. Females and males were given a choice between several odour sources that included (1) clean air, (2) uninfested, (3) host-infested, and (4) mechanically damaged cucumber plants. Females and males showed different preferences for volatiles emanating from these plants. Females responded significantly longer to the volatiles from uninfested plants than clean air, and to host-infested plants than uninfested plants. There were no significant differences in female responses to the volatiles from mechanically damaged and uninfested plants. Males responded significantly longer to clean air rather than uninfested plants. The volatiles from both uninfested and host-infested cucumber plants may play important roles in host habitat location of A. taragamae females.     

木虱啮小蜂对枸杞、枸杞木虱的行为反应

木虱啮小蜂Tetrastichus sp.是枸杞木虱Paratrioza sinica Yang & Li若虫外寄生蜂,是控制枸杞木虱种群数量动态的重要因子之一。为查明其寻找寄主的信息素,应用四臂嗅觉仪、培养皿两种方法测定了枸杞Lycium barbarum L.的健康叶、虫伤叶、枸杞木虱若虫、若虫与叶复合体、卵与叶复合体和若虫分泌物等不同物质对木虱啮小蜂行为反应的影响。结果证明枸杞木虱若虫分泌物对木虱啮小蜂有明显的吸引作用,四臂嗅觉仪测定时有73%的雌蜂趋向分泌物材料区（P=0.0000）。枸杞健康叶, 虫伤叶及其水、正丁烷提取物, 卵与枸杞叶复合体对木虱啮小蜂无明显吸引作用。木虱若虫水提物及正丁烷提取物在培养皿中可吸引木虱啮小蜂并可引起刺探行为,但嗅觉仪测定时单独若虫并不吸引木虱啮小蜂,只有活体若虫与枸杞叶片复合体才对木虱啮小蜂有吸引作用（P=0.0004）。木虱啮小蜂通过与寄主若虫的接触,可提高其搜索效率。接触过寄主若虫的雌蜂第一次找到寄主的时间显著少于未曾接触寄主若虫的雌蜂,前者只用后者1/4的时间,说明该寄生蜂能对其寄主若虫进行学习。     

The effect of repellents Ocimum forskolei and deet on the response of Anopheles stephensi to host odours

The behavioural response of Anopheles stephensi Liston (Diptera: Culicidae) to incubated host odours (from human and goat) and to human odour in combination with a repellent plant, Ocimum forskolei (Labiatae), or deet (N, N, diethyl-toluamide) (20%) was tested in a dual-port olfactometer. An. stephensi was significantly attracted to both host odours compared with air alone, but showed no preference when given a choice between both host odours simultaneously. In choice tests, the addition of deet to human odour did not significantly divert mosquitoes to human odour alone, but did divert them to goat odour. O. forskolei combined with human odour diverted mosquitoes to goat or human odour alone. Combinations of human odour and O. forskolei, and human odour and deet were both as unattractive as air alone, and attracted mosquitoes equally when offered simultaneously. The results indicate that use of O. forskolei as a repellent would be beneficial in reducing vector biting if used in communities in areas with partially zoophilic mosquito species such as An. stephensi, and where animals are present.     

The influence of host plant species on parasitism of pollen beetles (Meligethes spp.) by Phradis morionellus

We investigated the response of the specialist insect predator Oligota kashmirica benefica (Coleoptera: Staphylinidae) to volatiles from lima bean leaves infested with the spider mite Tetranychus urticae (Acari: Tetranychidae), both in a Y-tube olfactometer and in a field in Kyoto, Japan. Adult male and female predators were significantly more attracted to T. urticae-infested leaves than to clean air. Adult male and female predators were not more attracted to uninfested leaves, artificially damaged leaves, or the spider mites and their visible products when compared to clean air. In a field trap experiment, 12 adult predators were caught in three traps containing T. urticae-infested lima bean plants over 13 days, whereas no adult predators were trapped in three traps containing uninfested lima bean plants during the same period. These results showed that O. kashmirica benefica adults responded to herbivore-induced plant volatiles from T. urticae-infested lima bean leaves under both laboratory and field conditions.     

Role of volatiles emitted by host and non-host plants in the foraging behaviour of Dentichasmias busseolae, a pupal parasitoid of the spotted stemborer Chilo partellus

The role of volatiles from stemborer host and non‐host plants in the host‐finding process of Dentichasmias busseolae Heinrich (Hymenoptera: Ichneumonidae) a pupal parasitoid of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) was studied. The non‐host plant, molasses grass (Melinis minutiflora Beauv. (Poaceae)), is reported to produce some volatile compounds known to be attractive to some parasitoid species. The studies were conducted to explore the possibility of intercropping stemborer host plants with molasses grass in order to enhance the foraging activity of D. busseolae in such a diversified agro‐ecosystem. Olfactometric bioassays showed that volatiles from the host plants maize, Zea mays L., and sorghum, Sorghum bicolor (L.) (Poaceae), were attractive to the parasitoid. Infested host plants were the most attractive. Volatiles from molasses grass were repellent to the parasitoid. Further tests showed that volatiles from infested and uninfested host plants alone were preferred over those from infested and uninfested host plants combined with the non‐host plant, molasses grass. In dual choice tests, the parasitoid did not discriminate between volatiles from maize infested by either of the two herbivore species, C. partellus or Busseola fusca Fuller (Lepidoptera: Noctuidae). Volatiles from sorghum infested by C. partellus were preferred over those from C. partellus‐infested maize. The study showed that the pupal parasitoid D. busseolae uses plant volatiles during foraging, with those from the plant–herbivore complex being the most attractive. The fact that volatiles from molasses grass were deterrent to the parasitoid suggested that intercropping maize or sorghum with molasses grass was not likely to enhance the foraging behaviour of D. busseolae. Volatiles from the molasses grass may hinder D. busseolae's host location efficiency.     

Reactions of adult female parasitoids, particularly Aphidius rhopalosiphi, to volatile chemical cues from the host plants of their aphid prey

Abstract. In Y-tube olfactometer tests, Aphidius ervi Hal., Trioxys sp., Praon sp., Aphelinus flavus (Nees), Lysiphlebus fabarum (Marsh.) and Aphidius rophalosiphi De Stef. responded positively to the odour of the plant on which aphid mummies containing them had been collected. The response to host plant odour was greater than the response to the odour of host aphids, their honeydew or a combination of the two. The strongest response was to a combination of plant and host aphids. A. rhopalosiphi showed a strong positive response to three wheat volatiles (cis-3-hexenyl acetate, cis-3-hexen-1-ol and trans-2-hexenal) as well as to indole-3-acetaldehyde (a breakdown product of tryptophan in aphid honeydew). In both olfactometer tests with odours and choice trials with whole plants, newly emerged A. rhopalosiphi distinguished and preferred the variety of wheat on which their development had occurred to other wheat varieties.     

Host race formation inTetranychus urticae: genetic differentiation, host plant preference, and mate choice in a tomato and a cucumber strain

The two-spotted spider mite,Tetranychus urticae Koch, occurs in two colour forms in greenhouses in the Netherlands: a red form on tomato and a green form on cucumber. The evolutionary status of these strains was analysed by studying genetic differentiation, host plant preference, and mate choice. Males of the tomato strain preferred the female (30 h-old) teleiochrysales from the same strain to those of the cucumber strain, independent of the host plants (tomato, cucumber, bean) on which the teleiochrysales were placed. In contrast, males of the cucumber strain were not selective. In a Y-tube olfactometer, females of the cucumber strain were not responsive to host plant volatiles alone. However, in two-choice disc experiments, where females were exposed to both volatile and contact cues, they settled on cucumber leaves in preference to tomato leaves. Females of the tomato strain preferred the odour of tomato leaves and settled on tomato leaves in preference to cucumber leaves. These experimental results provide the first evidence for (1) host-plant independent mate selection in male spider mites and (2) olfactory discrimination between host plants in female spider mites. Electrophoretic analysis showed much genetic differentiation at the phosphoglucose isomerase locus. The cucumber strain showed large variation with 5 alleles, whereas the tomato strain was fixed for the most common allele of the cucumber strain. The results suggest that the two strains represent host races. We hypothesize that the tomato strain has originated from the cucumber strain because (1) tomato represents a more hostile host plant to spider mites (due to toxic compounds and glandular hairs) and (2) the tomato strain is genetically impoverished suggesting that it passed through one or more bottlenecks.